,,dragons ,,of ,,the ,,air              
,illustration3 ,,fig4 #dg4              
,,rhamphorhynchus ,,phyllunus           
,,showing ,,the ,,preservation ,,of     
,,the ,,wing ,,membranes      ,from the 
,lithographic slate of ,eichstaedt1     
,bavaria                                
,frontispiece                           
,,dragons ,,of ,,the ,,air              
,,an ,,account ,,of                     
,,extinct ,,flying ,,reptiles           
,,by                           ,h4 ,g4  
,,seeley1 ,f4,r4,s4    ,,professor ,,of 
,,geology ,,in ,,king',s ,,college1     
,,london2 ,,lecturer ,,on ,,geology     
,,and ,,mineralogy ,,in ,,the ,,royal   
,,indian ,,engineering ,,college        
,,with ,,eighty ,,illustrations         
,i ,,am ,a ,,brother ,,of ,,dragons     
,job xxx4 #bi                           
,,new ,,york3 ,d4 ,,appleton  ,,co4     
,,london3 ,,methuen  ,,co4              
#aija     ,,preface   ,i was a student  
of law at a time when ,sir ,richard     
,owen was lecturing on ,extinct ,fossil 

,reptiles4 ,the skill of the great      
master1 who built bones together as a   
child builds with a box of bricks1      
taught me that the laws which determine 
the forms of animals were less          
understood at that time than the laws   
which govern the relations of men in    
their country4 ,the laws of ,nature     
promised a better return of new         
knowledge for reasonable study4 ,a      
lecture on ,flying ,reptiles determined 
me to attempt to fathom the mysteries   
which gave new types of life to the     
,earth and afterwards took them away4   
,thus ,i became the very humble servant 
of the ,dragons of the ,air4 ,knowing   
but little about them ,i went to        
,cambridge1 and for ten years worked    
with the ,professor of ,geology1 the    
late ,rev4 ,adam ,sedgwick1 ,,ll4,d41   
,f4,r4,s41 in gathering their bones from
 the so-called ,cambridge ,coprolite    
bed1 the ,cambridge ,greensand4 ,the    
bones came in thousands1 battered and   
broken1 but instructive as better       

materials might not have been4 ,my rooms
 became filled with remains of existing 
birds1 lizards1 and mammals1 which threw
 light on the astonishing collection of 
old bones which ,i assisted in bringing 
together for the ,university4  ,in time 
,i had something to say about ,flying   
,animals which was new4 ,the story was  
told in the theatre of the ,royal       
,institution1 in a series of lectures4  
,some of them were repeated in several  
,english towns4 ,there was still much to
 learn of foreign forms of flying       
animals2 but at last1 with the aid of   
the ,government grant administered by   
the ,royal ,society1 and the chiefs of  
the great ,continental museums1 ,i saw  
all the specimens in ,europe4  ,so ,i   
have again written out my lectures1 with
 the aid of the latest discoveries1 and 
the story of animal structure has lost  
nothing in interest as a twice-told     
tale4 ,it still presents in epitome the 
story of life on the ,earth4 ,he who    
understands whence the ,flying ,reptiles

 came1 how they endured1 and disappeared
 from the ,earth1 has solved some of the
 greatest mysteries of life4 ,i have    
only contributed something towards      
solving the problems4  ,in telling my   
story1 chiefly of facts in ,nature1 an  
attempt is made to show how a naturalist
 does his work1 in the hope that perhaps
 a few readers will find happiness in   
following the workings of the laws of   
life4 ,such an illumination has proved  
to many worth seeking1 a solid return   
for labour1 which is not to be marketed 
on the ,exchange1 but may be taken      
freely without exhausting the treasury  
of ,nature's truths4 ,such outlines of  
knowledge as here are offered to a      
larger public1 may also1 ,i believe1 be 
acceptable to students of science and   
scientific men4  ,the drawings given in 
illustration of the text have been made 
for me by ,miss ,e4 ,b4 ,seeley4        
,h4 ,g4 ,s4   ,,kensington1 ,may1 #aija 
,,contents                              
,,page   ,,chapter ,i4   ,,flying       

,,reptiles                              
#a    ,,chapter ,,ii4   ,,how ,a        
,,reptile ,,is ,,known                  
#d    ,,chapter ,,iii4   ,a ,,reptile   
,,is ,,known ,,by ,,its ,,bones         
#aa    ,,chapter ,,iv4   ,,animals      
,,which ,,fly                           
#ae    ,,chapter ,v4   ,,discovery ,,of 
,,the ,,pterodactyle                    
#bg    ,,chapter ,,vi4   ,,how ,,animals
 ,,are ,,interpreted ,,by ,,their       
,,bones                          #cg    
,,chapter ,,vii4   ,,interpretation ,,of
 ,,pterodactyles ,,by ,,their ,,soft    
,,parts                 #de    ,,chapter
 ,,viii4   ,,the ,,plan ,,of ,,the      
,,skeleton                              
#eh    ,,chapter ,,ix4   ,,the          
,,backbone1 ,,or ,,vertebral ,,column   
#gh    ,,chapter ,x4   ,,the            
,,hip-,,girdle ,,and ,,hind ,,limb      
#ic    ,,chapter ,,xi4                  
,,shoulder-,,girdle ,,and ,,fore ,,limb 
#ajg    ,,chapter ,,xii4   ,,evidences  
,,of ,,the ,,animal',s ,,habits ,,from  

,,its ,,remains                  #acd   
,,chapter ,,xiii4   ,,ancient           
,,ornithosaurs ,,from ,,the ,,lias      
#adc    ,,chapter ,,xiv4                
,,ornithosaurs ,,from ,,the ,,middle    
,,secondary ,,rocks                     
#aec    ,,chapter ,,xv4   ,,ornithosaurs
 ,,from ,,the ,,upper ,,secondary       
,,rocks                        #agb     
,,chapter ,,xvi4   ,,classification ,,of
 ,,the ,,ornithosauria                  
#ahg    ,,chapter ,,xvii4   ,,family    
,,relations ,,of ,,pterodactyles ,,to   
,,animals ,,which ,,lived ,,with ,,them 
#aif    ,,chapter ,,xviii4   ,,how      
,,pterodactyles ,,may ,,have            
,,originated                            
#bac    ,,appendix                      
#bca    ,,index                         
#bcc     ,,list ,,of ,,illustrations    
,,fig4                                  
,,page   #dg4 ,wings of ,rhamphorhynchus
                                        
,frontispiece    #a4 ,lung of the       
lung-fish ,ceratodus                    

#e    #b4 ,attachment of the lower jaw  
in a ,mammal and in a ,pterodactyle     
#ab    #c4 ,chaldaean ,dragon           
#ae    #d4 ,winged human figure from the
 ,temple of ,ephesus                    
#af    #e4 ,flying fish ,exocoetus      
#ah    #f4 ,flying ,frog                
#ai    #g4 ,flying ,lizard 7,draco7     
#bj    #h4 ,birds in flight             
#bb    #i4 ,flying ,squirrel 7,pteromys7
                                        
#bd   #aj4 ,bats1 flying and walking    
#be   #aa4 ,skeleton of ,pterodactylus  
longirostris                        #bh 
#ab4 ,the skeleton restored             
#bi   #ac4 ,the animal form restored    
#cj   #ad4 ,fore limbs in four types of 
mammals                             #ch 
#ae4 ,pneumatic foramen in ,pterodactyle
 bone                          #df      
#af4 ,lungs of the bird ,apteryx        
#dh   #ag4 ,air cells in the body of an 
,ostrich                             #di
   #ah4 ,lung of a ,chameleon           
#ea   #ai4 ,brain in ,pterodactyle1     

,mammal1 ,bird1 and ,reptiles           
#ec   #bj4 ,skull of ,kingfisher and    
,rhamphorhynchus                        
#fc   #ba4 ,skull of ,heron and         
,rhamphorhynchus                        
#fe   #bb4 ,palate of ,macrocercus and 8
 ,campylognathus                        
#ga   #bc4 ,lower jaw of ,echidna and   
,ornithostoma                           
#gf   #bd4 ,first two neck vertebrae of 
,ornithocheirus                         
#ha   #be4 ,middle neck vertebrae of    
,ornithocheirus                         
#hc   #bf4 ,back vertebra of            
,ornithocheirus and ,crocodile          
#hf   #bg4 ,sacrum1 with hip bones1 of  
,rhamphorhynchus                        
#hh   #bh4 ,extremity of tail of        
,rhamphorhynchus phyllurus              
#ia   #bi4 ,hip-girdle bones in ,apteryx
 and ,rhamphorhynchus                   
#ie   #cj4 ,pelvis with prepubic bone in
 ,pterodactylus                      #if
   #ca4 ,pelvis with prepubic bones in  
,rhamphorhynchus                   #ig  

#cb4 ,pelvis of an ,alligator seen from 
below                          #ih      
#cc4 ,femora3 ,echidna1 ,ornithocheirus1
 ,ursus                         #ajj    
#cd4 ,tibia and fibula3 ,dimorphodon and
 ,vulture                      #ajb     
#ce4 ,metatarsus and digits in three    
,pterodactyles                   #ajd   
#cf4 ,sternum in ,cormorant and         
,rhamphorhynchus                        
#ajh   #cg4 ,sternum in ,ornithocheirus 
#aji   #ch4 ,shoulder-girdle bones in a 
bird and three ,pterodactyles           
#aac   #ci4 ,the ,notarium from the back
 of ,ornithocheirus                     
#aae   #dj4 ,the shoulder-girdle of     
,ornithocheirus                         
#aae   #da4 ,humerus of ,pigeon and     
,ornithocheirus                         
#aai   #db4 ,fore-arm of ,golden ,eagle 
and ,dimorphodon                        
#abj   #dc4 ,wrist bones of             
,ornithocheirus                         
#abd   #dd4 ,clawed digits of the hand  
in two ,pterodactyles                   

#abe   #de4 ,claw from the hand of      
,ornithocheirus                         
#abi   #df4 ,the hand in ,archaeopteryx 
and the ,ostrich                        
#acj   #dh4 ,slab of ,lias with bones of
 ,dimorphodon          ,to face page    
#adc   #di4 ,dimorphodon 7restored form7
 at rest                            #add
   #ej4 ,dimorphodon 7restored form of  
the animal7       ,to face page #ade    
#ea4 ,dimorphodon skeleton1 walking as a
 quadruped                 #adf   #eb4  
,dimorphodon skeleton as a biped        
#adg   #ec4 ,lower jaw of ,dorygnathus  
#adi   #ed4 ,dimorphodon 7wing membranes
 spread for flight7  ,to face page #aej 
#ee4 ,pelvis of ,dimorphodon            
#aea   #ef4 ,rhamphorhynchus skeleton   
7restored7                              
#afa   #eg4 ,scaphognathus 7restoration 
of #ahge7                               
#afc   #eh4 ,six restorations of        
,ornithosaurs                           
#afd   #ei4 ,ptenodracon skeleton       
7restored7                              

#afg   #fj4 ,cycnorhamphus suevicus slab
 with bones        ,to face page #afh   
#fa4 ,cycnorhamphus suevicus 7form of   
the animal7   ,to face page #afi   #fb4 
,cycnorhamphus suevicus skeleton        
7restored7                   #agj   #fc4
 ,cycnorhamphus ,fraasi 7restored       
skeleton form           of the animal7  
,to face page #agj   #fd4 ,cycnorhamphus
 ,fraasi 7restoration of the form       
of the body7                            
,to face page #aga   #fe4 ,neck vertebra
 of ,doratorhynchus from the ,purbeck   
#agc   #ff4 ,neck bone of ,ornithodesmus
 from the ,wealden                      
#agc   #fg4 ,sternum of ,ornithodesmus1 
seen from the front                     
#age   #fh4 ,sternum of ,ornithodesmus1 
side view1 showing the keel             
#age   #fi4 ,diagram of known parts of  
skull of ,ornithocheirus                
#agg   #gj4 ,neck bone of               
,ornithocheirus                         
#agi   #ga4 ,jaws of ,ornithocheirus    
from the ,chalk                         

#ahj   #gb4 ,palate of the ,english     
,toothless ,pterodactyle                
#aha   #gc4 ,two views of the skull of  
,ornithostoma 7,pteranodon7             
#ahb   #gd4 ,skeleton of ,ornithostoma  
#ahc   #ge4 ,comparison of six skulls of
 ,ornithosaurs                          
#aib   #gf4 ,pelvis of ,ornithostoma    
#aie   #gg4 ,skull of ,anchisaurus and  
,dimorphodon                            
#aii   #gh4 ,skull of ,ornithosuchus and
 ,dimorphodon                           
#bja   #gi4 ,the pelvis in ,ornithosaur 
and ,dinosaur                           
#bjd   #hj4 ,the prepubic bones in      
,dimorphodon and ,iguanodon             
#bjf      ,these figures are greatly    
reduced in size1 and when two or more   
bones are shown in the same figure all  
are brought to the same size     to     
facilitate the comparison4     ,,dragons
 ,,of ,,the ,,air     ,,chapter ,i      
,,flying ,,reptiles   ,the history of   
life on the earth during the epochs of  
geological time unfolds no more         

wonderful discovery among types of      
animals which have become extinct than  
the family of fossils known as flying   
reptiles4 ,its coming into existence1   
its structure1 and passing away from the
 living world are among the great       
mysteries of ,nature4  ,the animals are 
astonishing in their plan of            
construction4 ,in aspect they are unlike
 birds and beasts which1 in this age1   
hover over land and sea4 ,they gather   
into themselves in the body of a single 
individual1 structures which1 at the    
present day1 are among the most         
distinctive characters of certain       
mammals1 birds1 and reptiles4  ,the name
 flying reptile expresses this anomaly4 
,its invention is due to the genius of  
the great ,french naturalist ,cuvier1   
who was the first to realise that this  
extinct animal1 entombed in slabs of    
stone1 is one of the wonders of the     
world4  ,the word reptile has impressed 
the imagination with unpleasant sound1  
even when the habits of the animals it  

indicates are unknown4 ,it is familiarly
 associated with life which is reputed  
venomous1 and is creeping and cold4 ,its
 common type1 the serpent1 in many parts
 of the world takes a yearly toll of    
victims from man and beast1 and has     
become the representative of silent1    
active strength1 dreaded craft1 and     
danger4  ,science uses the word reptile 
in a more exact way1 to define the      
assemblage of cold-blooded animals which
 in familiar description are separately 
named serpents1 lizards1 turtles1       
hatteria1 and crocodiles4  ,turtles and 
the rest of them survive from great     
geological antiquity4 ,they present from
 age to age diversity of aspect and     
habit1 and in unexpected differences of 
outward proportion of the body show how 
the laws of life have preserved each    
animal type4 ,for the vital organs which
 constitute each animal a reptile1 and  
the distinctive bony structures with    
which they are associated1 remain       
unaffected1 or but little modified1 by  

the animal's external change in         
appearance4  ,the creeping reptile is   
commonly imagined as the antithesis of  
the bird4 ,for the bird overcomes the   
forces that hold even man to the earth1 
and enjoys exalted aerial conditions of 
life4 ,therefore the marvel is shared   
equally by learned and unlearned1 that  
the power of flight should have been an 
endowment of animals sprung from the    
breed of serpents1 or crocodiles1       
enabling them to move through the air as
 though they too were of a heaven-born  
race4 ,the wonder would not be lessened 
if the animal were a degraded           
representative of a nobler type1 or if  
it should be demonstrated that even     
beasts have advanced in the battle of   
life4 ,the winged reptile1 when compared
 with a bird1 is not less astounding    
than the poetic conceptions in ,milton's
 ,paradise ,lost of degradation which   
overtakes life that once was amongst the
 highest4 ,and on the other hand1 from  
the point of view of the teaching of    

,darwin in the theories of modern       
science1 we are led to ask whether a    
flying reptile may not be evidence of   
the physical exaltation which raises    
animals in the scale of organisation4   
,the dominance upon the earth of flying 
reptiles during the great middle period 
of geological history will long engage  
the interest of those who can realise   
the complexity of its structure1 or care
 to unravel the meaning of the          
procession of animal forms in successive
 geological ages which preceded the     
coming of man4  ,the outer vesture of an
 animal counts for little in estimating 
the value of ties which bind orders of  
animals together1 which are included in 
the larger classes of life4 ,the kindred
 relationship which makes the snake of  
the same class as the tortoise is       
determined by the soft vital            
organs--brain1 heart1 lungs--which are  
the essentials of an animal's existence 
and control its way of life4 ,the wonder
 which science weaves into the meaning  

of the word reptile1 bird1 or mammal1 is
 partly in exhibiting minor changes of  
character in those organs and other soft
 parts1 but far more in showing that    
while they endure unchanged1 the hard   
parts of the skeleton are modified in   
many ways4 ,for the bones of the reptile
 orders stretch their affinities in one 
direction towards the skeletons of      
salamanders and fishes2 and extend them 
also at the same time in other          
directions1 towards birds and mammals4  
,this mystery we may hope to partly     
unravel4     ,,chapter ,,ii  ,,how ,a   
,,reptile ,,is ,,known   ,,definition   
,,of ,,reptiles ,,by ,,their ,,vital    
,,organs  ,the relations of reptiles to 
other animals may be stated so as to    
make evident the characters and         
affinities which bind them together4    
,early in the nineteenth century        
naturalists included with the ,reptilia 
the tribe of salamanders and frogs which
 are named ,amphibia4 ,the two groups   
have been separated from each other     

because the young of ,amphibia pass     
through a tadpole stage of development4 
,they then breathe by gills1 like       
fishes1 taking oxygen from the air which
 is suspended in water1 before lungs are
 acquired which afterwards enable the   
animals to take oxygen directly from the
 air4 ,the amphibian sometimes sheds the
 gills1 and leaves the water to live on 
land4 ,sometimes gills and lungs are    
retained through life in the same       
individual4 ,this amphibian condition of
 lung and gill being present at the same
 time is paralleled by a few fishes     
which still exist1 like the ,australian 
,ceratodus1 the lung-fish1 an ancient   
type of fish which belongs to early days
 in geological time4  ,this             
metamorphosis has been held to separate 
the amphibian type from the reptile     
because no existing reptile develops    
gills or undergoes a metamorphosis4 ,yet
 the character may not be more important
 as a ground for classification than the
 community of gills and lungs in the    

fish and amphibian is ground for putting
 them together in one natural group4    
,for although no gills are found in     
reptiles1 birds1 or mammals1 the embryo 
of each in an early stage of development
 appears to possess gill-arches1 and    
gill-clefts between them1 through which 
gills might have been developed1 even in
 the higher vertebrates1 if the         
conditions of life had been favourable  
to such modification of structure4 ,in  
their bones ,reptiles and ,amphibia have
 much in common4 ,nearly all true       
reptiles lay eggs1 which are defined    
like those of birds by comparatively    
large size1 and are contained in shells4
 ,this condition is not usual in        
amphibians or fishes4 ,when hatched the 
young reptile is completely formed1 the 
image of its parent1 and has no need to 
grow a covering to its skin like some   
birds1 or shed its tail like some       
tadpoles4 ,the reptile is like the bird 
in freedom from important changes of    
form after the egg is hatched2 and the  

only structure shed by both is the      
little horn upon the nose1 with which   
the embryo breaks the shell and emerges 
a reptile or a bird1 growing to maturity
 with small subsequent variations in the
 proportions of the body4               
,illustration3 ,,fig4 #a  ,,lung ,,of   
,,the ,,fish ,,ceratodus    ,partly laid
 open to show its chambered structure   
7,after ,guenther7   ,,the ,,reptile    
,,skin  ,between one class of animals   
and another the differences in the      
condition of the skin are more or less  
distinctive4 ,in a few amphibians there 
are some bones in the skin on the under 
side of the body1 though the skin is    
usually naked1 and in frogs is said to  
transmit air to the blood1 so as to     
exercise a respiratory function of a    
minor kind4 ,this naked condition1 so   
unlike the armoured skin of the true    
,reptilia1 appears to have been         
paralleled by a number of extinct groups
 of fossils of the ,secondary rocks1    
such as ,ichthyosaurs and ,plesiosaurs1 

which were aquatic1 and probably also by
 some ,dinosauria1 which were           
terrestrial4  ,living reptiles are      
usually defended with some kind of      
protection to the skin4 ,among snakes   
and lizards the skin has commonly a     
covering of overlapping scales1 usually 
of horny or bony texture4 ,the tortoise 
and turtle tribe shut up the animal in a
 true box of bone1 which is cased with  
an armour of horny plates4 ,crocodiles  
have a thick skin embedding a less      
continuous coat of mail4 ,thus the skin 
of a reptile does not at first suggest  
anything which might become an organ of 
flight2 and its dermal appendages1 or   
scales1 may seem further removed from   
the feathers which ensure flying powers 
to the bird than from the naked skin of 
a frog4   ,,the ,,reptile ,,brain       
,although the mode of development of the
 young and the covering of the skin are 
conspicuous among important characters  
by which animals are classified1 the    
brain is an organ of some importance1   

although of greater weight in the higher
 ,vertebrata than in its lower groups4  
,reptiles have links in the mode of     
arrangement of the parts of their brains
 with fishes and amphibians4 ,the       
regions of that organ are commonly      
arranged in pairs of nervous masses1    
known as 7#a7 the olfactory lobes1 7#b7 
the cerebrum1 behind which is the minute
 pineal body1 followed by 7#c7 the pair 
of optic lobes1 and hindermost of all   
7#d7 the single mass termed the         
cerebellum4 ,these parts of the brain   
are extended in longitudinal order1 one 
behind the other in all three groups4   
,the olfactory lobes of the brain in    
,fishes may be as large as the cerebrum2
 but among ,reptiles and ,amphibians    
they are relatively smaller1 and they   
assume more of the condition found in   
mammals like the ,hare or ,mole1 being  
altogether subordinate in size4 ,and the
 cerebral masses begin to be wider and  
higher than the other parts of the      
brain1 though they do not extend forward

 above the olfactory lobes1 as is often 
seen in ,mammals4 ,in ,crocodiles the   
cerebral hemispheres have a tendency to 
a broad circular form4 ,among ,chelonian
 reptiles that region of the brain is   
more remarkable for height4 ,lizards and
 ,ophidians both have this part of the  
brain somewhat pear-shaped1 pointed in  
front1 and elongated4 ,the amphibian    
brain only differs from the lizard type 
in degree2 and differences between      
lizards' and amphibian brains are less  
noticeable than between the other orders
 of reptiles4 ,the reptilian brain is   
easily distinguished from that of all   
other animals by the position and       
proportions of its regions 7see ,fig4   
#ai1 p4 #ec74  ,birds have the parts of 
the brain formed and arranged in a way  
that is equally distinctive4 ,the       
cerebral lobes are relatively large and 
convex1 and deserve the descriptive name
 hemispheres4 ,they are always smooth1  
as among the lower ,mammals1 and extend 
backward so as to abut against the hind 

brain1 termed the cerebellum4 ,this     
junction is brought about in a peculiar 
way4 ,the cerebral hemispheres in a bird
 do not extend backward to override the 
optic lobes1 and hide them1 as occurs   
among adult mammals1 but they extend    
back between the optic lobes1 so as to  
force them apart and push them aside1   
downward and backward1 till they extend 
laterally beyond the junction of the    
cerebrum with the cerebellum4 ,the brain
 of a ,bird is never reptilian2 but in  
the young ,mammal the brain has a very  
reptilian aspect1 because both have     
their parts primarily arranged in a     
line4 ,therefore the brain appears to   
determine the boundary between bird and 
reptile exactly4   ,,reptilian          
,,breathing ,,organs  ,the breathing    
organs of ,birds and ,reptiles which are
 associated with these different types  
of brain are not quite the same4 ,the   
,frog has a cellular lung which1 in the 
details of the minute sacs which branch 
and cluster at the terminations of the  

tubes1 is not unlike the condition in a 
,mammal4 ,in a mammal respiration is    
aided by the bellows-like action of the 
muscles connected with the ribs1 which  
encase the cavity where the lungs are   
placed1 and this structure is absent in 
the ,frog and its allies4 ,the ,frog1 on
 the other hand1 has to swallow air in  
much the same way as man swallows water4
 ,the air is similarly grasped by the   
muscles1 and conveyed by them downward  
to the lungs4 ,therefore a ,frog keeps  
its mouth shut1 and the animal dies from
 want of air if its mouth is open for a 
few minutes4  ,crocodiles commonly lie  
in the sun with their mouths widely     
open4 ,the lungs in both ,crocodiles and
 ,turtles are moderately dense1         
traversed by great bronchial tubes1 but 
do not differ essentially in plan from  
those of a ,frog1 though the great      
branches of the bronchial tubes are     
stronger1 and the air chambers into     
which the lung is divided are somewhat  
smaller4 ,the ,new ,zealand ,hatteria   

has the lungs of this cellular type1    
though rather resembling the amphibian  
than the ,crocodile4 ,the lungs during  
life in all these animals attain        
considerable size1 the maximum          
dimensions being found in the           
terrestrial tortoises1 which owe much of
 their elevated bulk to the dimensions  
of the air cells which form the lungs4  
,the lungs of ,serpents and ,lizards are
 formed on a different plan4 ,in both   
those groups of reptiles the dense      
cellular tissue is limited to the part  
of the lung which is nearest to the     
throat4 ,this network of blood vessels  
and air cells extends about the         
principal bronchial tube much as in     
other animals1 but as it extends        
backward the blood vessels become few   
until the tubular lung appears in its   
hinder part1 as it extends down the     
body1 almost as simple in structure as  
the air bladder of a fish4 ,among       
,serpents only one of these tubular     
lungs is commonly present1 and the      

structure has a less efficient          
appearance as a breathing organ than the
 single lung of the fish ,ceratodus     
7,fig4 #a74 ,the ,chameleons are a group
 of lizards which differ in many ways   
from most of their nearest kindred1 and 
the lungs1 while conforming in general  
plan to the lizard type in being dense  
at the throat1 and a tubular bladder in 
the body1 give off on both sides a      
number of short lateral branches like   
the fingers of a glove 7,fig4 #ah1 p4   
#ea74  ,thus the breathing organs of    
reptiles present two or three distinct  
types which have caused ,serpents and   
,lizards to be associated in one group  
by most naturalists who have studied    
their anatomy2 while ,crocodiles and    
,chelonians represent a type of lung    
which is quite different1 and in those  
groups has much in common4 ,these       
characters of the breathing organs      
contribute to separate the cold-blooded 
armoured reptiles from the warm-blooded 
birds clothed with feathers1 as well as 

from the warm-blooded mammals which     
suckle their young2 for both these      
higher groups have denser and more      
elastic spongy lung tissue4  ,it will be
 seen hereafter that many birds in the  
most active development of their        
breathing organs substantially revert to
 the condition of the ,serpent or       
,chameleon in a somewhat modified way4  
,because1 instead of having one great   
bronchial tube expanded to form a vast  
reservoir of air which can be discharged
 from the lung in which the reptile has 
accumulated it1 the bird has the lateral
 branches of the bronchial tubes        
prolonged so as to pierce the walls of  
the lung1 when its covering membrane    
expands to form many air cells1 which   
fill much of the cavity of the bird's   
body 7see ,fig4 #af74 ,thus the bird    
appears to combine the characters of    
such a lung as that of a ,crocodile1    
with a condition which has some analogy 
with the lung of a ,chameleon4 ,it is   
this link of structure of the breathing 

organs between reptiles and birds that  
constitutes one of the chief interests  
of flying reptiles1 for they prove to   
have possessed air cells prolonged from 
the lungs1 which extended into the      
bones4     ,,chapter ,,iii  ,a ,,reptile
 ,,is ,,known ,,by ,,its ,,bones   ,such
 are a few illustrations of ways in     
which reptiles resemble other animals1  
and differ from them1 in the organs by  
means of which the classification of    
animals is made4 ,but such an idea is   
incomplete without noticing that the    
bony framework of the body associated   
with such vital organs also shows in its
 chief parts that reptiles are easily   
recognised by their bones4 ,i will      
therefore briefly state how reptiles are
 defined in some regions of the         
skeleton1 for in tracing the history of 
reptile life the bones are the principal
 remains of animals preserved in the    
rocks2 and the soft organs which have   
perished can only be inferred to have   
been present from the persistence of    

durable characteristic parts of the     
skeleton1 which are associated with     
those soft organs in animals which exist
 at the present day1 and are unknown in 
other animals in which the skeleton is  
different4   ,,the ,,hang ,,of ,,the    
,,lower ,,jaw  ,the manner in which the 
lower jaw is connected with the skull   
yields one of the most easily recognised
 differences between the great groups of
 vertebrate animals4  ,in ,mammals4--,in
 every mammal--such as the ,dog or      
,sheep--the lower jaw1 which is formed  
of one bone on each side1 joins directly
 on to the head of the animal1 and moves
 upon a bone of the skull which is named
 the temporal bone4 ,this character is  
sufficient to prove1 by the law of      
association of soft and hard parts of   
the body1 that such an animal had warm  
blood and suckled its young4            
,illustration3 ,,fig4 #b                
,,pterodactylus ,,kochi   ,,skull ,,of  
,,bear    ,comparison to show the       
articulation with the lower jaw in a    

mammal   and ,pterodactylus ,kochi4 ,the
 quadrate bone is lettered ,q in this   
,pterodactyle1 and comes between the    
skull and the lower jaw like the        
quadrate bone in a bird and in lizards4 
,in ,birds4--,in birds a great          
difference is found in this region of   
the head4 ,the temporal bone1 which it  
will be more convenient to name the     
squamosal bone1 from its squamous or    
scale-like form1 is still a part of the 
brain case1 and assists in covering the 
brain itself1 exactly as among mammals4 
,but the lower jaw is now made up of    
five or six bones4 ,and between the     
hindermost and the squamosal there is an
 intervening bar of bone1 unknown among 
mammalia1 which moves upon the skull by 
a joint1 just as the lower jaw moves    
upon it4 ,this movable bone unites with 
parts of the palate and the face1 and is
 known as the quadrate bone4 ,its       
presence proves that the animal         
possessing it laid eggs1 and if the face
 bones join its outer border just above 

the lower jaw1 it proves that the animal
 possessed hot blood4  ,in              
,reptiles4--,all reptiles are also      
regarded as possessing the quadrate     
bone4 ,but the squamosal bone with which
 it always unites is in less close union
 with the brain case1 and never covers  
the brain itself4 ,serpents show an     
extreme divergence in this condition    
from birds1 for the squamosal bone      
appears to be a loose external plate of 
bone which rests upon the compact brain 
case and gives attachment to the        
quadrate bone which is as free as in a  
bird4 ,among ,lizards the quadrate bone 
is usually almost as free4 ,in the other
 division of existing ,reptilia1        
including ,crocodiles1 the ,new ,zealand
 lizard-like reptile ,hatteria1 called  
,tuatera1 and ,turtles1 the squamosal   
and quadrate bones are firmly united    
with the bones of the brain case1 face1 
and palate1 so that the quadrate bone   
has no movement2 and the same condition 
appears in amphibians1 such as ,toads   

and ,frogs4 ,with these conditions of   
the quadrate bone are associated cold   
blood1 terrestrial life1 and young      
developed from eggs4  ,in               
,fishes4--,bony fishes1 and all others  
in which separate bones build up the    
skull1 differ from ,reptiles and ,birds 
much as those animals differ from       
,mammals4 ,the union of the lower jaw   
with the skull becomes complicated by   
the presence of additional bones4 ,the  
quadrate bone still forms a pulley      
articulation upon which the lower jaw   
works1 but between it and the squamosal 
bone is the characteristic bone of the  
fish known as the hyomandibular1        
commonly connected with opercular bones 
and metapterygoid which intervene1 and  
help to unite the quadrate with the     
brain case4 ,in the ,cartilaginous      
fishes there is only one bone connecting
 the jaws with the skull on each side4  
,this appears to prove that just as the 
structure of the arch of bones          
suspending the jaw may be complicated by

 the mysterious process called          
segmentation1 which separates a bone    
into portions1 so simplification and    
variation may result because the        
primitive divisions of the material     
cease to be made which exists before    
bones are formed4  ,the principal       
regions of the skull and skeleton all   
vary in the chief groups of animals with
 backbones2 so that the ,reptile may be 
recognised among fossils1 even in       
extinct groups of animals and           
occasionally restored from a fragment1  
to the aspect which characterised it    
while it lived4     ,,chapter ,,iv      
,,animals ,,which ,,fly   ,the nature of
 a reptile is now sufficiently          
intelligible for something to be said   
concerning flight1 and structures by    
means of which some animals lift        
themselves in the air4 ,it is not       
without interest to remember that1 from 
the earliest periods in human records1  
representations have been made of       
animals which were furnished with wings1

 yet walked upon four feet1 and in their
 typical aspect have the head shaped    
like that of a bird4 ,they are commonly 
named ,dragons4   ,,flying ,,dragons    
,illustration3 ,,fig4 #c  ,from ,the    
,battle between ,bel and the ,dragon    
,the effigy of the dragon survives to   
the present day in the figure over which
 ,st4 ,george triumphs1 on the reverse  
of the ,british sovereign4 ,in the      
luxuriant imaginations of ancient       
,eastern peoples1 dating back to        
prehistoric ages1 perhaps #ejjj ,b4,c41 
the dragons present an astonishing      
constancy of form4 ,in after-times they 
underwent a curious evolution1 as the   
conception of ,babylon and ,egypt is    
traced through ,assyria to ,greece4 ,the
 ,wings1 which had been associated at   
first with the fore limb of the typical 
dragon1 become characteristic of the    
,lion1 and of the poet's winged ,horse1 
and finally of the ,human figure itself1
 carved on the great columns of the     
,greek temples of ,ephesus4 ,these      

flying animals are historically         
descendants of the same common stock    
with the dragons of ,china and ,japan1  
which still preserve the aspect of      
reptiles4 ,their interest is chiefly in 
evidence of a latent spirit of evolution
 in days too remote for its meaning to  
be now understood1 which has carried the
 winged forms higher and ever higher in 
grade of organisation1 till their wings 
ceased to be associated with feelings of
 terror4 ,the ,hebrew cherubim are      
regarded by ,h4 ,e4 ,ryle1 ,bishop of   
,exeter1 as probably ,dragons1 and the  
figure of the conventional angel is the 
human form of the ,dragon4              
,illustration3 ,,fig4 #d4  ,,figure     
,,from ,,the ,,temple ,,of ,,ephesus    
,,organs ,,of ,,flight  ,turning from   
this reference to the realm of mythology
 to existing nature1 the power of flight
 is popularly associated with all the   
chief types of vertebrate               
animals--fishes1 frogs1 lizards1 birds1 
and mammals4 ,many of the animals ill   

deserve the name of flyers1 and most are
 exceptions to different conditions of  
existence which control their kindred1  
but it is convenient to examine for a   
little the nature of the structures by  
which this movement in the air1 which is
 not always flight1 is made possible4   
,certain fishes1 like the lung-fish     
,ceratodus1 of ,queensland1 and the     
mud-fish ,lepidosiren1 are capable of   
leaving the water and living on land1   
and for a time breathe air4 ,but neither
 these fishes nor ,periophthalmus1 which
 runs with rapid movement of its fins   
and carries the body more or less out of
 water1 or the climbing perch1 ,anabas1 
carried out of water over the country by
 ,indian jugglers1 ever put on the      
slightest approach to wings4   ,,flying 
,,fishes    ,illustration3 ,,fig4 #e4   
,,the ,,flying ,,fish ,,exocoetus       
,with the fins extended moving through  
the air  ,the flight of fishes is a kind
 of parachute support not unlike that by
 which a folded paper is made to travel 

in the air4 ,it is chiefly seen in the  
numerous species of a genus ,exocoetus1 
allied to the gar-pike 7,belone71 which 
is common in tropical seas1 and usually 
from a foot to eighteen inches long4    
,they emerge from the water1 and for a  
time support themselves in the air by   
means of the greatly developed breast   
fins1 which sometimes extend backward to
 the tail fin4 ,although these fins     
appear to correspond to the fore limbs  
of other animals1 they may not be moved 
at the will of the fish like the wing of
 a bird4 ,when the flying fishes are    
seen in shoals in the vicinity of ships1
 those fins remain extended1 so that the
 fish is said sometimes to travel #bjj  
yards at a speed of fifteen miles an    
hour1 rising twenty feet or more above  
the surface of the sea1 travelling in a 
straight line1 though sometimes         
influenced by the wind4 ,here the organ1
 which is at once a fin and a wing1     
consists of a number of thin long rods1 
or rays1 which are connected by         

membrane1 and vary in length to form an 
outline not unlike the wing of a bird   
which tapers to a point4 ,the interest  
of these animals is chiefly in the fact 
that flight is separated from the       
condition of having lungs with which it 
is associated in birds1 for although the
 flying fish has an air bladder1 there  
is no duct to connect it with the       
throat4   ,,flying ,,frogs              
,illustration3 ,,fig4 #f4  ,,the        
,,flying ,,frog 7,,rhacophorus7    ,the 
membranes of the foot and hand extend   
between the metatarsal and   metacarpal 
bones1 as well as the bones of the      
digits4  ,among amphibians the organs of
 flight are also of a parachute kind1   
but of a different nature4 ,they are    
seen in certain frogs which frequent    
trees1 and are limited to membranes     
which extend between the diverging      
digits of the hand and foot1 forming    
webs as fully developed as in the foot  
of a swimming bird4 ,as these frogs     
leap1 the membranes are expanded and    

help to support the weight of the body1 
so that the animal descends more easily 
as it moves from branch to branch4      
,there is no evidence that the bones of 
the digits ever became elongated like   
the fin rays of the flying fish or the  
wing bones of a ,bat2 but the web       
suggests the basis of such a wing1 and  
the possibilities under which wings may 
first originate1 by elongation of the   
bones of a webbed hand like that of a   
,flying ,frog4   ,,flying ,,lizards     
,illustration3 ,,fig4 #g4  ,,the        
,,flying ,,dragon1 ,,draco    ,forming a
 parachute by means of the extended ribs
  ,the ,reptilia in their several orders
 are remarkable for absence of any      
modification of the arms which might    
suggest a capacity for acquiring wings1 
as being latent in their organisation4  
,crocodiles1 ,tortoises1 and ,serpents  
are alike of the earth1 and not of the  
air4 ,but among ,lizards there are small
 groups of animals in which a limited   
capacity for movement through the air is

 developed4 ,it is best known in the    
family of small lizards named ,dragons1 
represented typically by the species    
,draco volans found in the ,oriental    
region of the ,east ,indies and ,malay  
,archipelago4  ,the organ of flight is  
produced in an unexpected way1 by means 
of the ribs instead of the limbs4 ,the  
ribs extend outward as far as the arms  
can stretch1 and the first five or six  
are prolonged beyond the body so as to  
spread a fold of skin on each side      
between the arm and the leg4 ,the       
membrane admits of some movement with   
the ribs4 ,this arrangement forms a     
parachute1 which enables the animal to  
move rapidly among branches of trees1   
extending the structure at will1 so that
 it is used with rapidity too quick to  
be followed by the eye1 as it leaps     
through considerable distances4  ,a less
 singular aid to movement in the air is 
found in some of the lizards termed     
,geckos4 ,the so-called ,flying ,gecko  
7,platydactylus homalocephalus7 has a   

fringe unconnected with ribs1 which     
extends laterally on the sides of the   
body and tail1 as well as at the back   
and front of the fore and hind limbs1   
and between the digits1 where the web is
 sometimes almost as well developed as  
among ,tree ,frogs4 ,this is essentially
 a lateral horizontal frill1 extending  
round the body4 ,its chief interest is  
in the circumstance that it includes a  
membrane which extends between the wrist
 bones and the shoulder on the front of 
the arm4 ,that is the only part of the  
fringe which represents the wing        
membrane of a bird4 ,the fossil flying  
reptiles have not only that membrane1   
but the lateral membranes at the sides  
of the body and behind the arms4  ,other
 lizards have the skin developed in the 
direction of the circumference of the   
body4 ,in the ,australian               
,chlamydosaurus it forms an immense     
frill round the neck like a mediaeval   
collar4 ,but though such an adornment   
might break a fall1 it could not be     

regarded as an organ of flight4         
,,flying ,,birds    ,illustration3      
,,fig4 #h4  ,,position ,,of ,,birds ,,in
 ,,flight  ,the wings of birds1 when    
they are developed so as to minister to 
flight1 are all made upon one plan2 but 
as examples of the variation which the  
organs contributing to make the fore    
limb manifest1 ,i may instance the short
 swimming limb of the ,penguin1 the     
practically useless rudiment of a wing  
found in the ,ostrich or ,kiwi1 and the 
fully developed wing of the ,pigeon4    
,the wings of birds obtain an extensive 
surface to support the animal by        
muscular movements of three             
modifications of structure4 ,first1 the 
bones of the fore limb are so shaped    
that they cannot1 in existing birds1 be 
applied to the ground for support and be
 used like the limbs of quadrupeds1 and 
are therefore folded up at the sides of 
the body1 and carried in an unused or   
useless state so long as the animal hops
 on the ground or walks1 balancing its  

weight on the hind legs4 ,secondly1     
there are two small folds of skin1 less 
conspicuous than those on the arms of   
,geckos2 one is between the wrist bones 
and the shoulder1 and the smaller hinder
 membrane is between the upper arm and  
the body4 ,these membranous expansions  
are insignificant1 and would in         
themselves be inadequate to support the 
body or materially assist its movements4
 ,thirdly1 the bird develops appendages 
to the skin which are familiarly known  
as feathers1 and the large feathers     
which make the wing are attached to the 
skin covering the lower arm bone named  
the ulna1 and the other bones which     
represent the wrist and hand4 ,the area 
and form of the bird's wing are due to  
individual appendages to the skin1 which
 are unknown in any other group of      
animals4 ,between the extended wing of  
the ,albatross1 measuring eleven feet in
 spread1 and the condition in the ,kiwi 
of ,new ,zealand1 in which the wing is  
vanishing1 there is every possible      

variation in size and form4 ,as a rule1 
the larger the animal the smaller is the
 wing area4 ,the problem of the origin  
of the bird's wing is not to be         
explained by study of existing animals2 
for the rowing organ of the ,penguin1   
which in itself would never suggest     
flight1 becomes an organ of flight in   
other birds by the growth upon it of    
suitable feathers4 ,anyone who has seen 
the birds named ,divers feeding under   
water1 swimming rapidly with their      
wings1 might never suspect that they    
were also organs of aerial flight4 ,the 
,ostrich is even more interesting1 for  
it has not developed flight1 and still  
retains at the extremities of two of the
 digits the slender claws of a limb     
which was originally no wing at all1 but
 the support of a four-footed animal    
7,fig4 #df1 p4 #acj74   ,,flying        
,,mammals  ,flight is also developed    
among mammals4 ,the ,insectivora include
 several interesting examples of animals
 which are capable of a certain motion  

through the air4 ,in the tropical       
forests of the ,malay ,archipelago are  
animals known as ,flying ,squirrels1    
,flying ,opossums1 ,flying ,lemurs1     
,flying ,foxes1 in which the skin       
extends outward laterally from the sides
 of the body so as to connect the fore  
limbs with the hind limbs1 and is also  
prolonged backward from the hind limbs  
to the tail4 ,the four digits are never 
elongated2 the bones of the fore limb   
are neither longer nor larger than those
 of the hind limb1 and the foot         
terminates in five little claws as in   
other four-footed animals4 ,this        
condition is adapted for the arboreal   
life which those animals live1 leaping  
from branch to branch1 feeding on fruits
 and leaves1 and in some cases upon     
insects4 ,these mammals may be compared 
with the ,flying ,geckos among reptiles 
in their parachute-like support by      
extension of the skin1 which gives them 
one of the conditions of support which  
contribute to constitute flight4        

,illustration3 ,,fig4 #i4  ,,flying     
,,squirrel 7,,pteromys7  ,bats4--,one   
entire order of mammals--the ,bats--not 
only possess true wings1 but are capable
 of flight which is sustained1 and in   
some cases powerful4 ,the wings are     
clothed with short hair like the rest of
 the body1 and thus the instrument of   
flight is unlike that of a bird4 ,the   
flight of a ,bat differs from that of   
all other animals in being dependent    
upon a modification of the bones of the 
fore limb1 which1 without interfering   
with the animal's movements as a        
quadruped1 secures an extension of the  
wing which is not inferior in area to   
that which the bird obtains by          
elongation of the bones of the arm and  
fore-arm and its feathers4 ,the         
distinctive peculiarity of the ,bat's   
wing is in the circumstance that four of
 the digits of the hand have their bones
 prolonged to a length which is often   
equal to the combined length of the arm 
and fore-arm4 ,the bones of the digits  

diverge like the ribs of an umbrella1   
and between them is the wing membrane1  
which extends from the sides of the body
 outward1 unites the fore limb with the 
hind limb1 and is prolonged down the    
tail as in the ,flying ,foxes4 ,bats    
have a small membrane in front of the   
bones of the arm and fore-arm stretching
 between the shoulder and the wrist1    
which corresponds with the wing membrane
 of a bird2 but the remainder of the    
membranes in ,bats' wings are absent in 
birds1 because their function is        
performed by feathers which give the    
wing its area4 ,the elongated digits of 
the ,bat's wing are folded together and 
carried at the sides of the body as     
though they were a few quill pens       
attached to its wrist1 where the one    
digit1 which is applied to the ground in
 walking1 terminates in a claw4         
,illustration3 ,,fig4 #aj  ,,new        
,,zealand ,,bat ,,flying4 ,,barbastelle 
,,walking  ,the organs which support    
animals in the air are thus seen to be  

more or less dissimilar in each of the  
great groups of animals4 ,they fall into
 three chief types3 first1 the          
parachute2 secondly1 the wing due to the
 feathers appended to the skin2 and     
thirdly1 the wing formed of membrane1   
supported by enormous elongation of the 
small bones of the back of the hand and 
fingers4 ,the two types of true wings   
are limited to birds and bats2 and no   
living reptile approximates to          
developing such an organ of flight as a 
wing4 ,judged1 therefore1 by the method 
of comparing the anatomical structures  
of one animal with another1 which is    
termed comparative anatomy1 the         
existence of flying reptiles might be   
pronounced impossible4 ,but in the light
 which the revelations of geology       
afford1 our convictions become tempered 
with modesty2 and we learn that with    
,nature nothing is impossible in        
development of animal structure4        
,,chapter ,v  ,,discovery ,,of ,,the    
,,pterodactyle   ,late in the eighteenth

 century1 in #aghd1 a small fossil      
animal with wings began to be known     
through the writings of ,collini1 as    
found in the white lithographic         
limestone of ,solenhofen in ,bavaria1   
and was regarded by him as a former     
inhabitant of the sea4 ,the foremost    
naturalist of the time1 the citizen     
,cuvier--for it was in the days of the  
,french ,republic--in #ahja1 in lucid   
language1 interpreted the animal as a   
genus of ,saurians4 ,that word1 so      
familiar at the present day1 was used in
 the first half of the century to       
include ,lizards and ,crocodiles2 and   
described animals akin to reptiles which
 were manifestly related neither to     
,serpents nor ,turtles4 ,but the term   
saurian is no longer in favour1 and has 
faded from science1 and is interesting  
only in ancient history of progress4    
,the lizards soon became classed in     
close alliance with snakes4 ,and the    
crocodiles1 with the ,hatteria1 were    
united with chelonians4 ,most modern    

naturalists who use the term saurian    
still make it an equivalent of lizard1  
or an animal of the lizard kind4        
,,cuvier    ,illustration3 ,,fig4 #aa4  
,,pterodactylus ,,longirostris 7,cuvier7
    ,the remains are preserved with the 
neck arched over the back1 and the      
jaws opened upward  ,cuvier defined this
 fossil from ,solenhofen as             
distinguished by the extreme elongation 
of the fourth digit of the hand1 and    
from that character invented for the    
animal the name ,pterodactyle4 ,he tells
 us that its flight was not due to      
prolongation of the ribs1 as among the  
living lizards named ,dragons2 or to a  
wing formed without the digits being    
distinguishable from each other1 as     
among ,birds2 nor with only one digit   
free from the wing1 as among ,bats2 but 
by having the wing supported mainly by a
 single greatly elongated digit1 while  
all the others are short and terminate  
in claws4 ,cuvier described the amazing 
animal in detail1 part by part2 and such

 has been the influence of his clear    
words and fame as a great anatomist that
 nearly every writer in after-years1 in 
,french and in ,english1 repeated       
,cuvier's conclusion1 maintained to the 
end1 that the animal is a saurian4      
,illustration3 ,,fig4 #ab4  ,,the       
,,skeleton ,,of ,,pterodactylus         
,,longirostris    ,reconstructed from   
the scattered bones in ,fig4 #ad1       
showing the limbs   on the left side    
,long before fashion determined1 as an  
article of educated belief1 that fossil 
animals exist chiefly to bridge over the
 gaps between those which still survive1
 the scientific men of ,germany were    
inclined to see in the ,pterodactyle    
such an intermediate type of life4 ,at  
first ,soemmerring and ,wagler would    
have placed the ,pterodactyle between   
mammals and birds4   ,,goldfuss         
,illustration3 ,,fig4 #ac4  ,,the       
,,pterodactylus ,,longirostris          
,,restored   ,,from ,,the ,,remains ,,in
 ,,fig4 #aa    ,showing positions of the

 wing membranes with the animal at rest 
,but the accomplished naturalist        
,goldfuss1 who described another fine   
skeleton of a ,pterodactyle in #ahca1   
saw in this flying animal an indication 
of the course taken by ,nature in       
changing the reptilian organisation to  
that of birds and mammals4 ,it is the   
first flash of light on a dark problem1 
and its brilliance of inference has     
never been equalled4 ,its effects were  
seen when ,prince ,charles ,bonaparte1  
the eminent ornithologist1 in ,italy1   
suggested for the group the name        
,ornithosauria2 when the profound       
anatomist de ,blainville1 in ,france1   
placed the short-tailed animal in a     
class between ,reptiles and ,birds named
 ,pterodactylia2 and ,andreas ,wagner1  
of ,munich1 who had more ,pterodactyles 
to judge from than his predecessors1 saw
 in the fossil animal a saurian in      
transition to a bird4   ,,von ,,meyer   
,but the ,german interpretation is not  
uniform1 and ,hermann von ,meyer1 the   

banker-naturalist of ,frankfurt a4_/,m41
 who made himself conversant with all   
that his predecessors knew1 and enlarged
 knowledge of the ,pterodactyles on the 
most critical facts of structure1       
continued to regard them as true        
reptiles1 but flying reptiles4 ,such is 
the influence of von ,meyer that all    
parts of the world have shown a         
disposition to reflect his opinions1    
especially as they practically coincide 
with the earlier teaching of ,cuvier4   
,owen and ,huxley in ,england1 ,cope and
 ,marsh in ,america1 ,gaudry in ,france1
 and ,zittel in ,germany have all placed
 the ,pterodactyles as flying reptiles4 
,their judgment is emphatic4 ,but there 
is weight of competent opinion to       
endorse the evolutionary teaching of    
,goldfuss that they rise above reptiles4
 ,to form an independent opinion the    
modern student must examine the animals1
 weigh their characters bone by bone1   
familiarise himself1 if possible1 with  
some of the rocks in which they are     

found2 to comprehend the conditions     
under which the fossils are preserved1  
which have added not a little to the    
interest in ,pterodactyles1 and to the  
difficulty of interpretation4           
,,geological ,,history ,,of             
,,pterodactyles ,,in ,,germany  ,we may 
briefly recapitulate the geological     
history4 ,those remains of ,ornithosaurs
 which have been mentioned1 with a      
multitude of others which are the glory 
of the museums of ,munich1 ,stuttgart1  
,tuebingen1 ,heidelberg1 ,bonn1         
,haarlem1 and ,london1 have all been    
found in working the lithographic stone 
of ,bavaria4 ,the whitish yellow        
limestone forms low1 flat-topped hills1 
now isolated from each other by natural 
denudation1 which has removed the       
intervening rock4 ,the stone is found at
 some distance north of the ,danube1 in 
a line due north of ,augsburg1 in the   
country about ,pappenheim1 and          
especially at the villages of           
,solenhofen1 ,eichstaedt1 ,kelheim1 and 

,nusplingen4 ,these beds belong to the  
rocks which are named ,white ,jura      
limestone in ,germany1 which is of about
 the same geological age as the         
,kimeridge clay in ,england4 ,much of it
 divides into very thin layers1 and in  
these planes of separation the fossils  
are found4 ,they include the ,ammonites 
lithographicus and a multitude of marine
 shells1 king crabs and other           
,crustacea1 sea-urchins1 and other      
fossils1 showing that the deposit was   
formed in the sea4 ,the preservation of 
jelly-fish1 which so soon disappear when
 left dry on the beach1 shows that the  
ancient calcareous mud had unusual power
 of preserving fossils4 ,into this sea1 
with its fishes great and small1 came   
land plants from off the land1          
dragonflies and other insects1 tortoises
 and lizards1 ,pterodactyles with their 
flying organs1 and birds still clothed  
with feathers4 ,sometimes the wing      
membranes of the flying reptiles are    
found fully stretched by the wing       

finger1 as in examples to be seen at    
,munich and in the ,yale ,museum in     
,newhaven1 in ,america4 ,at ,haarlem    
there is an example in which the wing   
membrane appears to be folded much as in
 the wing of a ,bat1 when the animal    
hangs suspended1 with the flying        
membrane bent into a few wide           
undulations4  ,the ,solenhofen ,slate   
belongs to about the middle period of   
the history of flying reptiles1 for they
 range through the ,secondary epochs of 
geological time4 ,remains are recorded  
in ,germany from the ,keuper beds at the
 top of the ,trias1 which is the bottom 
division of the ,secondary strata2 and  
,i believe ,i have seen fragments of    
their bones from the somewhat older     
,muschelkalk of ,germany4   ,,their     
,,history ,,in ,,england  ,in ,england  
the remains are found for the first time
 in the ,lower ,lias of ,lyme ,regis1 in
 ,dorset1 and the ,upper ,lias of       
,whitby1 in ,yorkshire4 ,in ,wuertemberg
 they occur on the same horizons4 ,they 

reappear in ,england1 in every          
subsequent age1 when the conditions of  
the strata and their fossils give       
evidence of near proximity to land4 ,in 
the ,stonesfield ,slate of ,stonesfield1
 in ,oxfordshire1 the bones are found   
isolated1 but indicate animals of some  
size1 though not so large as the rare   
bones of reputed true birds which appear
 to have left their remains in the same 
deposit4  ,at least two ,pterodactyles  
are found in the ,oxford clay1 known    
from more or less fragmentary remains or
 isolated bones2 just as they occur in  
the ,kimeridge ,clay1 ,purbeck          
,limestone1 ,wealden sandstones1 and    
especially in newer ,secondary rocks1   
named ,gault1 ,upper ,greensand1 and    
,chalk1 in the south-east of ,england4  
,owing to exceptional facilities for    
collecting1 in consequence of the       
,cambridge ,greensand being excavated   
for the valuable mineral phosphate of   
lime it contains1 more than a thousand  
bones are preserved1 more or less broken

 and battered1 in the ,woodwardian      
,museum of the ,university of ,cambridge
 alone4 ,to give some idea of their     
abundance1 it may be stated that they   
were mostly gathered during two or three
 years1 as a matter of business1 by an  
intelligent foreman of washers of the   
nodules of phosphate of lime1 which1 in 
commerce1 are named coprolites4 ,he soon
 learned to distinguish ,pterodactyle   
bones from other fossils by their       
texture1 and learned the anatomical     
names of bones from specimens in the    
,university ,museum4 ,this workman1 ,mr4
 ,pond1 employed by ,mr4 ,william       
,farren1 brought together not only the  
best of the remains at ,cambridge1 but  
most of those in the museums at ,york   
and in ,london1 and the thousands of    
less perfect specimens in public and    
private collections which passed through
 the present writer's hands in          
endeavours to secure for the ,university
 useful illustrations of the animal's   
structure4 ,these fragments1 among which

 there are few entire bones1 are        
valuable1 for they have afforded        
opportunities of examining the articular
 ends of bones in every aspect1 which is
 not possible when similar organic      
remains are embedded in rock in their   
natural connexions4  ,in ,england       
,flying ,reptiles disappear with the    
,chalk4 ,in that period they were widely
 distributed1 being found in ,bohemia1  
in ,brazil1 and ,kansas in the ,united  
,states1 as well as in ,kent and other  
parts of ,england4 ,they attained their 
largest dimensions in this period of    
geological time4 ,one imperfect fragment
 of a bone from the ,laramie rocks of   
,canada was described1 ,i believe1 by   
,cope1 though not identified by him as  
,ornithosaurian1 and is probably newer  
than other remains4   ,,aspect ,,of     
,,pterodactyles  ,if this series of     
animals could all be brought together   
they would vary greatly in aspect and   
stature1 as well as in structure4 ,some 
have the head enormously long1 in others

 it is large and deep1 characters which 
are shared by extinct reptiles which do 
not fly1 and to which some birds may    
approximate2 while in a few the head is 
small and compact1 no more conspicuous1 
relatively1 than the head of a ,sparrow4
 ,the neck may be slender like that of a
 ,heron1 or strong like that of an      
,eagle2 the back is always short1 and   
the tail may be inconspicuous1 or as    
long as the back and neck together4     
,these flying reptiles frequently have  
the proportions of the limbs similar to 
those of a ,bat1 with fore legs strong  
and hind legs relatively small2 while in
 some the limbs are as long1            
proportionately1 and graceful as those  
of a ,deer4 ,with these differences in  
proportions of the body are associated  
great differences in the relative length
 of the wing and spread of the wing     
membranes4   ,,dimensions ,,of ,,the    
,,animals  ,the dimensions of the       
animals have probably varied in all     
periods of geological time4 ,the        

smallest1 in the ,lithographic ,slate1  
are smaller than ,sparrows1 while       
associated with them are others in which
 the drumstick bone of the leg is eight 
inches long4 ,in the ,cambridge         
,greensand and ,chalk imperfect         
specimens occur1 showing that the upper 
arm bones are larger than those of an   
,ox4 ,the shaft is one and a half inches
 in diameter and the ends three inches  
wide4 ,such remains may indicate        
,pterodactyles not inferior in size to  
the extinct ,moas of ,new ,zealand1 but 
with immensely larger heads1 animals far
 larger than birds of flight4  ,the late
 ,sir ,richard ,owen1 on first seeing   
these fragmentary remains1 said the     
flying reptile with outstretched pinions
 must have appeared like the soaring    
,roc of ,arabian romance1 but with the  
features of leathern wings with crooked 
claws superinduced1 and gaping mouth    
with threatening teeth4 ,eventually we  
shall obtain more exact ideas of their  
aspect1 when the structures of the      

several regions of the body have been   
examined4 ,the great dimensions of the  
stretch of wing1 often computed at      
twenty feet in the larger examples1     
might lead to expectations of great     
weight of body1 if it were not known    
that an albatross1 with wings spreading 
eleven feet1 only weighs about seventeen
 pounds4     ,,chapter ,,vi  ,,how      
,,animals ,,are ,,interpreted ,,by      
,,their ,,bones   ,there is only one    
safe path which the naturalist may      
follow who would tell the story of the  
meaning and nature of an extinct type of
 animal life1 and that is to compare it 
as fully as possible in its several     
bones1 and as a whole1 with other       
animals1 especially with those which    
survive4 ,it is easy to fix the place in
 nature of living animals and determine 
their mutual relations to each other1   
because all the organs--vital as well as
 locomotive--are available for          
comparison4 ,on such evidence they are  
grouped together into the large         

divisions of ,beasts1 ,birds1 and       
,reptiles2 as well as placed in smaller 
divisions termed ,orders1 which are     
based upon less important modifications 
of fundamental structures4 ,all these   
characteristic organs have usually      
disappeared in the fossil4 ,hence a new 
method of study of the hard parts of the
 skeleton1 which alone are preserved1 is
 used in the endeavour to discover how  
the ,flying ,reptile or other extinct   
animal is to be classified1 and how it  
acquired its characters or came into    
existence4   ,,variations ,,of ,,bones  
,,among ,,mammalia    ,illustration3    
,,fig4 #ad4  ,,the ,,fore ,,limb ,,in   
,,four ,,types ,,of ,,mammals           
,comparison of the fore limb in mammals1
 showing variation of form   of the     
bones with function  ,resemblances and  
differences in the bones are easily     
over-estimated in importance as evidence
 of pedigree relationship4 ,the         
,mammalia show1 by means of such        
skeletons as are exhibited in any       

,natural ,history ,museum1 how small is 
the importance to be attached to even   
the existence of any group of bones in  
determining its grade of organisation4  
,the whole ,whale tribe suckle their    
young and conform to the distinctive    
characters in brain and lungs which mark
 them as being mammals4 ,but if there is
 one part of the skeleton more than     
another which distinguishes the         
,mammalia1 it is the girdle of bones at 
the hips which supports the hind limbs4 
,it is characterised by the bone named  
the ilium being uniformly directed      
forward4 ,yet in the ,whale tribe the   
hip-girdle and the hind limb which it   
usually supports are so faintly         
indicated as to be practically lost2    
while the fore limb becomes a paddle    
without distinction of digits1 and is   
therefore devoid of hoofs or claws1     
which are usual terminations of the     
extremities in mammals4 ,yet this       
swimming paddle1 with its ill-defined   
bones--sometimes astonishing in number1 

as well as in fewness of the finger     
bones--is represented by the burrowing  
fore limb of the ,mole1 which lives     
underground2 by the elongated hoofed    
legs of the ,giraffe1 which lives on    
plains2 and the extended arm and finger 
bones of the ,bat1 which are equally    
mammals with the ,whale4 ,from such     
comparison it is seen that no           
proportion1 or form1 or length1 or use  
of the bones of the limbs1 or even the  
presence of limbs1 is necessarily       
characteristic of a mammal4 ,no         
limitation can be placed upon the       
possible diversity of form or           
development of bones in unknown animals1
 when they are considered in the light  
of such experience of varied structural 
conditions in living members of a single
 class4  ,what is true for the limbs and
 the bony arches which support them is  
true for the backbone also1 for the     
ribs1 and to some extent for the skull4 
,the neck in the ,whale is shortened    
almost beyond recognition4 ,in the      

,giraffe the same seven vertebrae are   
elongated into a marvellous neck2 so    
that in the technical definition of a   
mammal both are said to have seven neck 
vertebrae4 ,yet exceptions show a       
capacity for variation4 ,one of the     
,sloths reduces the number to six1 while
 another has nine vertebrae in the neck2
 proving that there is no necessary     
difference between a mammal and a       
reptile when judged by a character which
 is typically so distinctive of mammals 
as the number of the neck bones4  ,the  
skull varies too1 though to a less      
extent4 ,the ,great ,ant-eater of ,south
 ,america is a mammal absolutely without
 teeth4 ,the ,porpoises have a simple   
unvarying row of conical teeth with     
single roots extending along the jaw4   
,and the dental armature of the jaws1   
and relative dimensions of the skull    
bones1 exhibit such diversity1 in       
evidence of what may be parted with or  
acquired1 that recognition of the many  
reptilian structures and bones in the   

skull of ,ornithorhynchus1 the          
,australian ,duckbill1 demonstrates that
 the difficulties in recognising an     
animal by its bones are real1 unless we 
can discover the ,animal ,type to which 
the bones belong2 and that there is very
 little in osteology which may not be   
lost without affecting an animal's grade
 of organisation4   ,,variation ,,in    
,,skin ,,covering ,,of ,,mammals  ,even 
the covering of the body varies in the  
same class1 or even order of animals1 so
 that the familiar growth on the skin is
 never its only possible covering4 ,the 
,indian ant-eater1 named ,manis1 which  
looks like a gigantic fir-cone1 the     
,armadillo1 which sheathes the body in  
rings of bone1 bearing only a scanty    
development of hair1 are examples of    
mammalian hair1 as singular as the      
quills of a ,porcupine1 the horn of a   
,rhinoceros1 or the growth of hair of   
varying length and stoutness on         
different parts of the body in various  
animals1 or the imperfect development of

 hair in the marine ,cetacea4 ,among    
living animals it is enough for         
practical purposes to say that a mammal 
is clothed with hair1 but in a fossil   
state the hair must usually be lost     
beyond recognition from its fineness and
 shortness of growth4   ,,variation ,,in
 ,,skin ,,covering ,,of ,,birds  ,no    
,class of living animals is more        
homogeneous than ,birds2 and            
well-preserved remains prove that1 at   
least as far back in time as the ,upper 
,oolites1 birds were clothed with       
feathers of essentially the same mode of
 growth and appearance as the feathers  
of living birds4 ,there may1 therefore1 
be no ground for assuming that the      
covering was ever different1 though some
 regions of the skin are free from      
feathers4 ,yet the variations from fine 
under-down to the scale-like feathers on
 the wings of a ,penguin1 or the great  
feathers in the wings of birds of       
flight1 or the double quill of the      
,ostrich group1 are calculated to yield 

dissimilar impressions in a fossil      
state1 even if the fine down would be   
preserved in any stratum4   ,,variation 
,,in ,,the ,,bones ,,of ,,birds         
,osteologically there is less variety in
 the skeleton of birds than in other    
great groups of animals4 ,the existing  
representatives do not exhaust its      
capability for modification4 ,the few   
specimens of birds hitherto found in the
 ,secondary strata have rudely removed  
many differences in the bones which     
separated living birds from reptiles2 so
 that if only the older fossil birds    
were known1 and the ,tertiary and living
 birds had not existed1 a bird might    
have been defined as an animal having   
its jaw armed with teeth1 instead of    
devoid of teeth2 with vertebrae cupped  
at both ends1 instead of with a         
saddle-shaped articulation which in     
front is concave from side from side1   
and convex from above downwards2 in     
which the bones of the hand are         
separate1 so that three digits          

terminating in claws can be applied to  
the ground1 instead of the metacarpal   
bones being united in a solid mass with 
clawless digits2 and in which the tail  
is elongated like the tail of a lizard4 
,yet the limits to variation are not to 
be formulated till ,nature has exhausted
 all her resources in efforts to        
preserve organic types by adapting them 
to changed circumstances4 ,birds may be 
regarded theoretically as equally       
capable with mammals of parting with    
almost every distinctive structure in   
the skeleton by which it is best known4 
,even the living frigate bird blends the
 early joints of the backbone into a    
compact mass like a sacrum4 ,the        
,penguin has a cup-and-ball articulation
 in the early dorsal vertebrae1 with the
 ball in front4 ,and the genus ,cypselus
 has the upper arm bone almost as broad 
as long1 unlike the bird type4 ,such    
examples prove that we are apt to accept
 the predominant structures in an animal
 type as though they were universal1 and

 forget that inferences based1 like     
those of early investigators1 on limited
 materials may be re-examined with      
advantage4   ,,variation ,,in ,,the     
,,bones ,,of ,,reptiles  ,the true      
,reptilia1 notwithstanding some strong  
resemblances to ,birds in technical     
characters of the skeleton1 display     
among their surviving representatives an
 astonishing diversity in the bony      
framework of the body1 exceeding that of
 the mammalia4 ,this unlooked-for       
capacity for varying the plan of        
construction of the skeleton is in      
harmony with the diversity of structure 
in groups of extinct animals to which   
the name reptiles has also been given4  
,the interval in form is so vast between
 ,serpent and ,tortoise1 and so         
considerable in structure of the        
skeleton between these and the several  
groups of ,lizards1 ,crocodiles1 and    
,hatteria1 that any other diversity     
could not be more surprising4 ,and the  
inference is reasonable that just as    

mammals live in the air1 in the sea1 on 
the earth1 and burrow under the earth1  
similar modes of existence might be     
expected for birds and reptiles1 though 
no bird is yet known to have put on the 
aspect of a fish1 and no reptiles have  
been discovered which roamed in herds   
like antelopes1 or lived in the air like
 birds or bats1 unless these fossil     
flying animals prove on examination to  
justify the name by which they are      
known4  ,comparative study of structure 
in this way demolishes the prejudice1   
born of experience of the life which now
 remains on earth1 that the ideas of    
,reptile and of ,flight are incongruous1
 and not to be combined in one animal4  
,the comparative study of the parts of  
animals does not leave the student in a 
chaos of possibilities1 but teaches us  
that organic structures1 which mark the 
grades of life1 have only a limited     
scope of change2 while ,nature flings   
away every part of the skeleton which is
 not vital1 or changes its form with    

altering circumstances of existence1    
enforced by revolutions of the ,earth's 
surface in geological time1 in her      
efforts to save organisms from          
extinction and pass the grade of life   
onward to a later age4  ,the bones are  
only of value to the naturalist as      
symbols1 inherited or acquired1 and vary
 in value as evidence of the nature and 
association of those vital organs which 
differentiate the great groups of the   
vertebrata4  ,these distinctive         
structures1 which separate ,mammals1    
,birds1 and ,reptiles1 are sometimes    
demonstrated by the impress of their    
existence left on the bones2 or         
sometimes they may be inferred from the 
characters of the skeleton as a whole4  
,,chapter ,,vii  ,,interpretation ,,of  
,,pterodactyles ,,by ,,their ,,soft     
,,parts   ,,the ,,organs ,,which ,,fix  
,,an ,,animal',s ,,place ,,in ,,nature  
,we shall endeavour to ascertain what   
marks of its grade of organisation the  
,pterodactyle has to show4 ,the organs  

which are capable of modifying the bones
 are probably limited to the kidneys1   
the brain1 and the organs of            
respiration4 ,it may be sufficient to   
examine the latter two4   ,,pneumatic   
,,foramina ,,in ,,pterodactyles         
,illustration3 ,,fig4 #ae4  ,,head ,,of 
,,the ,,humerus ,,of ,,the              
,,pterodactyle   ,,ornithocheirus       
,showing position of the pneumatic      
foramen on the ulnar side of the   bone 
as in a bird  ,hermann von ,meyer1 the  
historian of the ,ornithosaurs of the   
,lithographic ,slate1 as early as #ahcg 
described some ,pterodactyle bones from 
the ,lias of ,franconia1 which showed   
that air was admitted into the interior 
of the bones by apertures near their    
extremities1 which1 from this           
circumstance1 are known as pneumatic    
foramina4 ,he drew the inference1       
naturally enough1 that such a structure 
is absolute proof that the ,pterodactyle
 was a flying animal4 ,it was not quite 
the right form in which the conclusion  

should have been stated1 because the    
,ostrich and other birds which do not   
fly have the principal bones pneumatic4 
,afterwards1 in #ahei1 the larger bones 
which ,professor ,sedgwick1 of          
,cambridge1 transmitted to ,sir ,richard
 ,owen established this condition as    
characteristic of the ,flying ,reptiles 
of the ,cambridge ,greensand4 ,it was   
thus found as a distinctive structure of
 the bones both at the beginning and the
 close of the geological history of     
these animals4 ,von ,meyer remarks that 
the supposition readily follows that in 
the respiratory process there was some  
similarity between ,pterodactyles and   
,birds4 ,this cautious statement may    
perhaps be due to the circumstance that 
in many animals air cavities are        
developed in the skull without being    
connected with organs of respiration4   
,it is well known that the bulk of the  
,elephant's head is due to the brain    
cavity being protected with an envelope 
formed of large air cells4 ,small air   

cells are seen in the skulls of oxen1   
pigs1 and many other mammals1 as well as
 in the human forehead4 ,the head of a  
bird like the ,owl owes something of its
 imposing appearance to the way in which
 its mass is enlarged by the dense      
covering of air cells in the bones above
 the brain1 like that seen in some      
,cretaceous ,pterodactyles4 ,nor are the
 skulls of ,crocodiles or ,tortoises    
exceptions to the general rule that an  
animal's head bones may be pneumatic    
without implying a pneumatic            
prolongation of air from the lungs4 ,the
 mere presence of air cells without     
specification of the region of the      
skeleton in which they occur is not     
remarkable4 ,the holes by which air     
enters the bones are usually much larger
 in ,pterodactyles than in ,birds1 but  
the entrance to the air cell prolonged  
into the bones is the same in form and  
position in both groups4 ,so far as can 
be judged by this character1 there is no
 difference between them4 ,the          

importance of the comparison can only be
 appreciated by examining the bones side
 by side4 ,in the upper arm bone of a   
bird1 on what is known as the ulnar     
border1 near to the shoulder joint1 and 
on the side nearest to it1 is the       
entrance to the air cell in the humerus4
 ,in the ,pterodactyle the corresponding
 foramen has the same position1 form1   
and size1 and is not one large hole1 but
 a reticulation of small perforations1  
one beyond another1 exactly such as are 
seen in the entrance to the air cell in 
the bone of a bird1 in which the        
pneumatic character is found4 ,for it is
 not every bird of flight which has this
 pneumatic condition of the bones2 and  
,dr4 ,crisp stated that quite a number  
of birds--the ,swallow1 ,martin1 ,snipe1
 ,canary1 ,wood-wren and ,willow-wren1  
,whinchat1 ,glossy-starling1            
,spotted-fly-catcher1 and ,black-headed 
,bunting--have no air in their bones4   
,and it is well known that in many      
birds1 especially water birds1 it is    

only the upper bones of the limbs which 
are pneumatic1 while the smaller bones  
retain the marrow4   ,,lungs ,,and ,,air
 ,,cells    ,illustration3 ,,fig4 #af4  
,,lungs ,,of ,,the ,,bird ,,apteryx     
,,partly ,,opened ,,on   ,,the          
,,right-,,hand ,,side    ,the circles   
are openings of the bronchial tubes on  
the surface of the   lung4 ,the notches 
on the inner edges of the lungs are     
impressions of   the ribs 7,after ,r4   
,owen7    ,illustration3 ,,fig4 #ag4    
,,the ,,body ,,of ,,an ,,ostrich ,,laid 
,,open ,,to ,,show ,,the   ,,air ,,cells
 ,,which ,,extend ,,through ,,its       
,,length 7,after ,georges ,roche7  ,it  
may be well to remember that the lungs  
of a bird are differently conditioned   
from those of any other animal4 ,instead
 of hanging freely suspended in the     
cone-shaped chamber of the thorax formed
 by the ribs and sternum1 they are      
firmly fixed on each side1 so that the  
ribs deeply indent them and hold them in
 place4 ,the lungs have the usual       

internal structure1 being made up of    
branching cells4 ,the chief peculiarity 
consists in the way in which the air    
passes not only into them1 but through  
them4 ,the air tube of the throat of a  
bird1 unlike that of a man1 has the     
organ of voice1 not at the upper end in 
the form of a larynx1 but at the lower  
end1 forming what is termed a syrinx4   
,there is no evidence of this in a      
fossil state1 although in a few birds   
the rings of the trachaea become        
ossified1 and are preserved4 ,but below 
the syrinx the trachaea divides into two
 bronchi1 tubes which carry the ringed  
character into the lungs for some       
distance1 and these give off branches   
termed bronchial tubes1 the finer       
subdivisions from which1 in their       
clustered minute branching sacs1 make up
 the substance of the lung4 ,there is   
nothing exceptional in that4 ,but       
towards the outer or middle part of the 
ventral or under surface of the lungs1  
four or five rounded openings are seen  

on each side4 ,each of these openings   
resembles the entrance of the air cell  
into a bone1 since it displays several  
smaller openings which lead to it4 ,each
 opening from the lung leads to an air  
cell4 ,those cells may be regarded as   
the blowing out of the membrane which   
covers the lungs into a film which holds
 air like a mass of soap bubbles1 until 
the whole cavity of the body of a bird  
from neck to tail is occupied by        
sacculated air cells1 commonly ten in   
number1 five on each side1 though two   
frequently blend at the base of the neck
 in the region of the ,v-shaped bone    
named the clavicle or furculum1         
popularly known as the merry-thought4   
,most people have seen some at least of 
these semi-transparent bladder-like air 
cells beneath the skin in the abdominal 
region of a fowl4 ,the cells have names 
from their positions1 and on each side  
one is abdominal1 two are thoracic1 one 
clavicular1 and one cervical1 which last
 is at the base of the neck4 ,the       

clavicular and abdominal air cells are  
perhaps the most interesting4 ,the air  
cell termed clavicular sends a process  
outward towards the arm1 along with the 
blood vessels which supply the arm4     
,thus this air cell1 entering the region
 of the axilla or arm-pit1 enters the   
upper arm bone usually on its under     
side1 close to the articular head of the
 humerus1 and in the same way the air   
may pass from bone to bone through every
 bone in the fore limb4 ,the hind limbs 
similarly receive air from the abdominal
 air cell1 which supplies the femur and 
other bones of the leg1 the sacrum1 and 
the tail4 ,but the joints of the        
backbone in front of the sacrum receive 
their air from the cervical air sac4    
,the air cells are not limited to the   
bones1 but ramify through the body1 and 
in some cases extend among the muscles4 
,a bird may be said to breathe not only 
with its lungs1 but with its whole body4
 ,and it is even affirmed that          
respiration has been carried on through 

a broken arm bone when the throat was   
closed1 and the bird under water4       
,birds differ greatly in the extent to  
which the aircell system prolonged from 
the lungs is developed1 some having the 
air absent from every bone1 while       
others1 like the ,swift1 are reputed to 
have air in every bone of the body4     
,comparison shows that in so far as the 
bones are the same in ,bird and         
,ornithosaur1 the evidence of the air   
cells entering them extends to          
resemblance1 if not coincidence1 in     
every detail4 ,no living group of       
animals except birds has pneumatic limb 
bones1 in relation to the lungs2 so that
 it is reasonable to conclude that the  
identical structures in the bones were  
due to the same cause in both the living
 and extinct groups of animals4 ,it is  
impossible to say that the lungs were   
identical in ,birds and ,pterodactyles1 
but so far as evidence goes1 there is no
 ground for supposing them to have been 
different4   ,,the ,,lungs ,,of         

,,reptiles    ,illustration3 ,,fig4 #ah4
  ,,the ,,side ,,of ,,the ,,body ,,of ,a
 ,,chameleon    ,ribs removed to show   
the sacculate branched form of the lung 
,there is nothing comparable to birds1  
either in the lungs of living reptiles  
or in their relation to the bones4 ,the 
,chameleon is remarkable in that the    
lung is not a simple bladder prolonged  
through the whole length of the body    
cavity1 as in a serpent1 but it develops
 a number of large lateral branches     
visible when the body is laid open4     
,except near the trachaea1 where the    
tissue has the usual density of a lizard
 lung1 the air cell is scarcely more    
complicated than the air bladder of a   
fish1 and does not enter into any bone  
of the skeleton4 ,and although many     
fishes like the ,loach have the swim    
bladder surrounded by bone connected    
with the head1 it offers no analogy to  
the pneumatic condition of the bones in 
the ,pterodactyle4   ,,the ,,form ,,of  
,,the ,,brain ,,cavity    ,illustration3

 ,,fig4 #ai4  ,,the ,,form ,,of ,,the   
,,brain  ,but the identity of the       
pneumatic foramina in ,birds and ,flying
 ,reptiles is not a character which     
stands by itself as evidence of         
organisation1 for a mould of the form of
 the brain case contributes evidence of 
another structural condition which      
throws some light on the nature of      
,ornithosaurs4 ,among many of the lower 
animals1 such as turtles1 the brain does
 not fill the chamber in the dry skull1 
in which the same bones are found as are
 moulded upon the brain in higher       
animals4 ,for the brain case in such    
reptiles is commonly an envelope of     
cartilage1 as among certain fishes2 and 
except among serpents1 the ,ophidia1 the
 bones do not completely close the      
reptilian brain case in front4 ,the     
brain fills the brain case completely   
among birds4 ,a mould from its interior 
is almost as definite in displaying the 
several parts of which it is formed as  
the actual brain would be4 ,and the     

chief regions of the brain in a         
bird--cerebrum1 optic lobes1            
cerebellum--show singularly little      
variation in proportion or position4    
,the essential fact in a bird's brain1  
which separates it absolutely from all  
other animals1 is that the pair of nerve
 masses known as the optic lobes are    
thrust out at the sides1 so that the    
large cerebral hemispheres extend partly
 over them as they extend between them  
to abut against the cerebellum4 ,this   
remarkable condition has no parallel    
among other vertebrate animals4 ,in     
,fishes1 ,amphibians1 ,reptiles1 and    
,mammals the linear succession of the   
several parts of the brain is never     
departed from2 and any appearance of    
variation from it among mammals is more 
apparent than real1 for the linear      
succession may be seen in the young calf
 till the cerebral hemispheres grow     
upward and lop backward1 so as to hide  
the relatively small brain masses which 
correspond to the optic lobes of        

reptiles1 extending over these          
corpora-quadrigemina1 as they are named1
 so as to cover more or less of the mass
 of the cerebellum4 ,from these         
conditions of the brain and skull1 it   
would not be possible to mistake a mould
 from the brain case of a bird for that 
of a reptile1 though in some conditions 
of preservation it is conceivable that  
the mould of the brain of a bird might  
be distinguished with difficulty from   
that of the brain in the lowest mammals4
 ,taken by itself1 the avian form of    
brain in an animal would be as good     
evidence that its grade of organisation 
was that of a bird as could be offered4 
,,the ,,brain ,,in ,,solenhofen         
,,pterodactyles  ,it happens that moulds
 of the brain of ,pterodactyles1 more or
 less complete1 are met with of all     
geological ages--,liassic1 ,oolitic1 and
 ,cretaceous4 ,the ,solenhofen ,slate is
 the only deposit in ,europe in which   
,pterodactyle skulls can be said to be  
fairly numerous4 ,they commonly have the

 bones so thin as to show the form of   
the upper surface of the mould of the   
brain1 or the bones have scaled off the 
mould1 or remain in the counterpart slab
 of stone1 so as to lay bare the shape  
of the brain mass4  ,in the ,museum at  
,heidelberg a skull of this kind is seen
 in the long-tailed genus of            
,pterodactyles named ,rhamphorhynchus4  
,it shows the large rounded cerebral    
hemispheres1 which extend in front of   
cerebral masses of smaller size a little
 below them in position1 which perhaps  
are as like the brain of a monotreme    
mammal as a bird4  ,the short-tailed    
,pterodactylus described by ,cuvier has 
the cerebral hemispheres very similar to
 those of a bird1 but the relations of  
the hinder parts of the brain to each   
other are less clear4  ,the first       
specimen to show the back of the brain  
was found by ,mr4 ,john ,francis        
,walker1 ,m4,a41 in the ,cambridge      
,greensand4 ,i was able to remove the   
thick covering of cellular bone which   

originally extended above it1 and thus  
expose evidence that in the mutual      
relations of the fore and hind parts of 
the brain bird and ornithosaur were     
practically identical4 ,another         
,cambridge ,greensand skull showed that 
in the genus ,ornithocheirus the optic  
lobes of the brain are developed        
laterally1 as in birds4 ,that skull was 
isolated and imperfect4 ,but about the  
same time the late ,rev4 ,w4 ,fox1 of   
,brixton1 in the ,isle of ,wight1       
obtained from ,wealden beds another     
skull1 with jaws1 teeth1 and the        
principal bones of the skeleton1 which  
showed that the ,wealden ,pterodactyle  
,ornithodesmus had a similar and        
bird-like brain4 ,in #ahhh ,mr4 ,e4 ,t4 
,newton1 ,f4,r4,s41 obtained a skull    
from the ,upper ,lias1 uncrushed and    
free from distortion4 ,this made known  
the natural mould of the brain1 which   
shows the cerebral hemispheres1 optic   
lobes1 and cerebellum more distinctly   
than in the specimens previously known4 

,in some respects it recalls the        
,heidelberg brain of ,rhamphorhynchus in
 the apparently transverse subdivision  
of the optic lobes1 but it is           
unmistakably bird-like1 and quite unlike
 any reptile4   ,,importance ,,of ,,the 
,,brain ,,and ,,breathing ,,organs  ,so 
far as the evidence goes1 it appears    
that these fossil flying animals show no
 substantial differences from birds1    
either in the mould of the brain or the 
impress of the breathing organs upon the
 bones4 ,these approximations to birds  
of the nervous and respiratory systems1 
which are beyond question two of the    
most important of the vital organs of an
 animal1 and distinctive beyond all     
others of birds1 place the naturalist in
 a singular dilemma4 ,he must elect     
whether he will trust his interpretation
 to the soft organs1 which among        
existing animals never vary their type  
in the great classes of vertebrate      
animals1 and on which the animal is     
defined as something distinct from its  

envelope the skeleton and its appendages
 the limbs1 or whether he will ignore   
them4 ,the answer must choose           
substantially between belief that the   
existing order of ,nature gives warrant 
for believing that these vital          
characteristics which have been         
discussed might equally coexist with the
 skeleton of a mammal or a reptile1 as  
with that of a bird1 for which there is 
no particle of evidence in existing     
life4 ,or1 as an alternative1 the fact  
must be accepted that birds only have   
such vital organs as are here found1 and
 therefore the skeleton1 that may be    
associated with them1 cannot affect the 
reference of the type to the same       
division of the animal kingdom as birds4
 ,the decision need not be made without 
further consideration4 ,but brain and   
breathing organs of the avian type are  
structures of a different order of      
stability in most animals from the      
bones1 which vary to a remarkable extent
 in almost every ordinal group of       

animals4   ,,temperature ,,of ,,the     
,,blood  ,the organs of circulation and 
digestion are necessarily unknown4      
,there are reasons why the blood may    
have been hot1 such as the evidences    
from the wings of exceptional activity2 
though the temperature depends more upon
 the amount of blood in the body than   
upon the apparatus by which it is       
distributed4 ,we speak of a ,crocodile  
as cold-blooded1 yet it is an animal    
with a four-chambered heart not         
incomparable with that of a bird4 ,on   
the other hand1 the ,tunny1 a sort of   
giant ,mackerel1 is a fish with a       
three-chambered heart1 only breathing   
the air dissolved in water1 which has   
blood as warm as a mammal1 its          
temperature being compared to that of a 
pig4 ,several fishes have blood as warm 
as that of ,manis1 the scaly ant-eater2 
and many birds have hotter blood than   
mammals4 ,the term hot-blooded1 as      
distinct from cold-blooded1 applied to  
animals1 is relative to the arbitrary   

human standard of experience1 and       
expresses no more than the circumstance 
that mammals and birds are warmer       
animals than reptiles and fishes4  ,the 
exceptional temperature of the ,flying  
,fish has led to a vague impression that
 physical activity and its effect upon  
the amount of blood which vigour of     
movement circulates1 are more important 
in raising an animal's temperature than 
possession of the circulatory organs    
commonly associated with hot blood1     
which drive the blood in distinct       
courses through the body and breathing  
organs4 ,yet the kind of heart which is 
always associated with vital structures 
such as ,pterodactyles are inferred to  
have possessed from the brain mould and 
the pneumatic foramina in the bones1 is 
the four-chambered heart of the bird and
 the mammal4 ,considering these organs  
alone--of which the fossil bones yield  
evidence--we might anticipate1 by the   
law of known association of structures1 
that nothing distinctly reptilian       

existed in the other soft part of the   
vital organisation1 because there is no 
evidence in favour of or against such a 
possibility4     ,,chapter ,,viii  ,,the
 ,,plan ,,of ,,the ,,skeleton   ,while  
these animals are incontestably nearer  
to birds than to any other animals in   
their plan of organisation1 thus far no 
proof has been found that they are      
birds1 or can be included in the same   
division of vertebrate life with        
feathered animals4 ,it is one of the    
oldest and soundest teachings of        
,linnaeus that a bird is known by its   
feathers2 and the record is a blank as  
to any covering to the skin in          
,pterodactyles4 ,there is the strongest 
probability against feathers having     
existed such as are known in the        
,archaeopteryx1 because every           
,solenhofen ,ornithosaur appears to have
 the body devoid of visible or          
preservable covering1 while the two     
birds known from the ,solenhofen ,slate 
deposit are well clothed with feathers  

in perfect preservation4 ,we turn from  
the skin to the skeleton4  ,the plan on 
which the skeleton is constructed       
remains as evidence of the animal's     
place in nature1 which is capable of    
affording demonstration on which        
absolute reliance would have been       
placed1 if the brain and pneumatic      
foramina had remained undiscovered4     
,with the entire skeleton before us1 it 
is inconceivable that anatomical science
 should fail to discover the true nature
 of the animal to which it belonged1 by 
the method of comparing one animal with 
another4 ,there is no lack of this kind 
of evidence of ,pterodactyles in the    
three or four scores of skeletons1 and  
thousands of isolated or associated     
bones1 preserved in the public museums  
of ,europe and ,america4  ,i may recall 
the circumstance that the discovery of  
skeletons of fossil animals has         
occasionally followed upon the          
interpretation of a single fragment1    
from which the animal has been well     

defined1 and sometimes accurately drawn1
 before it was ever seen4 ,so ,i        
propose1 before drawing any conclusions 
from the skeletons in the entirety of   
their construction1 to examine them bone
 by bone1 and region by region1 for     
evidence that will manifest the nature  
of this brood of ,dragons4 ,their living
 kindred1 and perhaps their extinct     
allies1 assembled as a jury1 may be able
 to determine whether resemblances exist
 between them1 and whether such         
similarity between the bones as exists  
is a common inheritance1 or is a common 
acquisition due to similar ways of life1
 and no evidence of the grade of the    
organism among vertebrate animals4  ,the
 bones of these ,ornithosaurs1 when     
found isolated1 first have to be        
separated from the organisms with which 
they are associated and mixed in the    
geological strata4 ,this discrimination 
is accomplished in the first instance by
 means of the texture of the surface4   
,the density and polish of the bones is 

even more marked than in the bones of   
birds1 and is usually associated with a 
peculiar thinness of substance of the   
bone1 which is comparable to the        
condition in a bird1 though usually a   
little stouter1 so that the bones resist
 crushing better4 ,pterodactyle bones in
 many instances are recognised by their 
straightness and comparatively uniform  
dimensions1 due to the exceptional      
number of long bones which enter into   
the structure of the wing as compared   
with birds4 ,when the bones are         
unerringly determined as                
,ornithosaurian1 they are placed side by
 side with all the bones which are most 
like them1 till1 judged by the standard 
of the structures of living animals1 the
 fossil is found to show a composite    
construction as though it were not one  
animal but many1 while its individual   
bones often show equally composite      
characters1 as though parts of the      
corresponding bone in several animals   
had been cunningly fitted together and  

moulded into shape4   ,,the ,,plan ,,of 
,,the ,,head ,,in ,,ornithosaurs  ,the  
head is always the most instructive part
 of an animal4 ,it is less than an inch 
long in the small ,solenhofen skeleton  
named ,pterodactylus brevirostris1 and  
is said to be three feet nine inches    
long in the toothless ,pterodactyle     
,ornithostoma from the ,chalk of        
,kansas4 ,most of these animals have a  
long1 slender1 conical form of head1    
tapering to the point like the beak of a
 ,heron1 forming a long triangle when   
seen from above or from the side4       
,sometimes the head is depressed in     
front1 with the beak flattened or       
rounded as in a ,duck or ,goose1 and    
occasionally in some ,wealden and       
,greensand species the jaws are         
truncated in front in a massive snout   
quite unlike any bird4 ,the back of the 
head is sometimes rounded as among      
birds1 showing a smooth pear-shaped     
posterior convexity in the region of the
 brain4 ,sometimes the back of the head 

is square and vertical or oblique4      
,occasionally a great crest of cellular 
tissue is extended backward from above  
the brain case over the spines of the   
neck bones4  ,there are always from two 
to four lateral openings in the skull4  
,first1 the nostril is nearest to the   
extremity of the beak4 ,secondly1 the   
orbits of the eyes are placed far       
backward4 ,these two openings are always
 present4 ,the nostril may incline      
upward4 ,the orbits of the eyes are     
usually lateral1 though their upper     
borders sometimes closely approximate1  
as in the woodpecker-like types from the
 ,solenhofen ,slate named ,pterodactylus
 ,kochi1 now separated as another genus4
 ,in most genera there is an opening in 
the side of the head1 between the eye   
hole and the nostril1 known as the      
antorbital vacuity2 and another opening1
 which is variable in size and known as 
the temporal vacuity1 is placed behind  
the eye4 ,the former is common in the   
skulls of birds1 the latter is absent   

from all birds and found in many        
reptiles4  ,the palate is usually       
imperfectly seen1 but ,english and      
,american specimens have shown that it  
has much in common with the palate in   
birds1 though it varies greatly in form 
of the bones in representatives from the
 ,lias1 ,oolites1 and ,cretaceous rocks4
  ,from the scientific aspect the       
relative size of the head1 its form1 and
 the positions and dimensions of its    
apertures and processes1 are of little  
importance in comparison with its plan  
of construction1 as evidenced by the    
positions and relations to each other of
 the bones of which it is formed4 ,there
 usually is some difficulty in stating  
the limits of the bones of the skull1   
because in ,pterodactyles1 as among     
birds1 they usually blend together1 so  
that in the adult animal the sutures    
between the bones are commonly          
obliterated4  ,bones have relations to  
each other and places in the head which 
can only change as the organs with which

 they are associated change their       
positions4 ,no matter what the position 
of a nostril may be--at the extremity of
 a long snout1 as in an ant-eater1 or   
far back at the top of the head in a    
porpoise1 or at the side of the head in 
a bird--it is always bordered by        
substantially the same bones1 which vary
 in length and size with the changing   
place of the nostril and the form of the
 head4 ,every region of the head is     
defined by this method of construction2 
so that eye holes and nose holes1 brain 
case and jaw bones1 palate and teeth1   
beak1 and back of the skull are all     
instructive to those who seek out the   
life-history of these animals4 ,we may  
briefly examine the head of an          
,ornithosaurian4   ,,bones ,,about ,,the
 ,,nostril  ,no matter what its form may
 be1 the head of an ,ornithosaur always 
terminates in front in a single bone    
called the intermaxillary4 ,it sends a  
bar of bone backward above the visible  
nostrils1 between them2 and a bar on    

each side forms the margin of the jaw in
 which teeth are implanted4 ,the bone   
varies in depth1 length1 sharpness1     
bluntness1 slenderness1 and massiveness4
 ,as the bone becomes long the jaw is   
compressed from side to side1 and the   
openings of the nostrils are removed    
backward to an increasing distance from 
the extremity of the beak4  ,the outer  
and hinder border of the nostril is made
 by another bone named the maxillary    
bone1 which is usually much shorter than
 the premaxillary4 ,it contains the     
hindermost teeth1 which rarely differ   
from those in front1 except in sometimes
 being smaller4  ,the nasal bones1 which
 always make the upper and hinder border
 of the nostrils1 meet each other above 
them1 in the middle line of the beak4   
,illustration3 ,,fig4 #bj    ,showing   
that the extremity of the jaws in       
,rhamphorhynchus was   sheathed in horn 
as in the giant ,kingfisher1 since the  
jaws   similarly gape in front4    ,the 
hyoid bones are below the lower jaw in  

the ,pterodactyle4  ,the nostrils are   
unusually large in the ,lias genus named
 ,dimorphodon1 and small in species of  
the genus ,rhamphorhynchus from         
,solenhofen4 ,such differences result   
from the relative dimensions and        
proportions of these three bones which  
margin the nasal vacuity1 and by varying
 growth of their front margins or of    
their hinder margins govern the form of 
the snout4  ,the jaws are most massive  
in the genera known from the ,wealden   
beds to the ,chalk4 ,the palatal surface
 is commonly flat or convex1 and often  
marked by an elevated median ridge which
 corresponds to a groove in the lower   
jaw1 though the median ridge sometimes  
divides the palate into two parallel    
concave channels4 ,the jaw is margined  
with teeth which are rarely fewer than  
ten or more than twenty on each side4   
,they are sharp1 compressed from side to
 side1 curved inward1 and never have a  
saw-like edge on the back and front     
margins4 ,no teeth occur upon the bones 

of the palate4  ,in most birds there is 
a large vacuity in the side of the head 
between the nostril and the orbit of the
 eye1 partly separated from it by the   
bone which carries the duct for tears   
named the lachrymal bone4 ,the same     
preorbital vacuity is present in all    
long-tailed ,pterodactyles1 though it is
 either less completely defined or      
absent in the group with short tails4   
,it affords excellent distinctive       
characters for defining the genera4 ,in 
the long-tailed genus ,scaphognathus    
from ,solenhofen this preorbital opening
 is much larger than the nostril1 while 
in ,dimorphodon these vacuities are of  
about equal size4 ,rhamphorhynchus is   
distinguished by the small size of the  
antorbital vacuity1 which is placed     
lower than the nostril on the side of   
the face4 ,the aperture is always       
imperfectly defined in ,pterodactylus1  
and is a relatively small vacuity       
compared with the long nostril4 ,in     
,ptenodracon the antorbital vacuity     

appears to have no existence separate   
from the nostril which adjoins the eye  
hole4 ,and so far as is known at present
 there is no lateral opening in advance 
of the eye in the skull in any          
,ornithosaur from ,cretaceous rocks1    
though the toothless ,ornithostoma is   
the only genus with the skull complete4 
,when a separate antorbital vacuity     
exists1 it is bordered by the maxillary 
bone in front1 and by the malar bone    
behind4 ,the prefrontal bone is at its  
upper angle4 ,that bone is known in a   
separate state in reptiles and1 ,i      
think1 in monotreme mammals4 ,its       
identity is soon lost in the mammal1 and
 its function in the skull is different 
from the corresponding bone in          
,pterodactyles4   ,,bones ,,about ,,the 
,,eyes    ,illustration3 ,,fig4 #ba4    
,,upper ,,surface ,,of ,,skull ,,of     
,,the ,,heron    ,compared with the same
 aspect of the skull of ,rhamphorhynchus
  ,the third opening in the side of the 
head1 counting from before backward1 is 

the orbit of the eye4 ,in this vacuity  
is often seen the sclerotic circle of   
overlapping bones formed in the external
 membrane of the eye1 like those in     
nocturnal birds and some reptiles4 ,the 
eye hole varies in form from an inverted
 pear-shape to an oblique or transverse 
oval1 or a nearly circular outline4 ,it 
is margined by the frontal bone above2  
the tear bone or lachrymal1 and the     
malar or cheek bone in front2 while the 
bones behind appear to be the           
quadrato-jugal and post-frontal bones1  
though the bones about the eye are      
somewhat differently arranged in        
different genera4  ,the eyes were       
frequently1 if not always1 in contact   
with the anterior walls of the brain    
case1 as in many birds1 and are always  
far back in the side of the head4 ,in   
,dimorphodon they are in front of the   
articulation of the lower jaw2 in       
,rhamphorhynchus1 above that            
articulation2 while in ,ornithostoma    
they are behind the articulation for the

 jaw4 ,this change is governed by the   
position of the quadrate bone1 which is 
vertical in the ,lias genus1 inclined   
obliquely forward in the fossils from   
the ,oolites1 and so much inclined in   
the ,chalk fossil that the small orbit  
is thrown relatively further back4      
,thus far the chief difference in the   
,pterodactyle skull from that of a bird 
is in the way in which the malar arch is
 prolonged backward on each side4 ,it is
 a slender bar of bone in birds1 without
 contributing ascending processes to    
border vacuities in the side of the     
face1 while in these fossil animals the 
lateral openings are partly separated by
 the ascending processes of these bones4
 ,this divergence from birds1 in the    
malar bone entering the orbit of the eye
 is approximated to among reptiles and  
mammals1 though the conditions1 and     
perhaps the presence of a bone like the 
post-orbital bone1 are paralleled only  
among ,reptiles4 ,the ,pterodactyles    
differ among themselves enough for the  

head to make a near approach to         
,reptiles in ,dimorphodon1 and to ,birds
 in ,pterodactylus4 ,in the ,ground     
,hornbill and the ,shoebill the         
lachrymal bones in front of the orbits  
of the eyes grow down to meet the malar 
bars without uniting with them4 ,the    
post-frontal region also is prolonged   
downward almost as far as the malar bar1
 as though to show that a bird might    
have its orbital circle formed in the   
same way and by the same bones as in    
,pterodactylus4 ,cretaceous             
,ornithosaurs sometimes differ from     
birds apparently in admitting the       
quadrato-jugal bone into the orbit4 ,it 
then becomes an expanded plate1 instead 
of a slender bar as in all birds4       
,,the ,,temporal ,,fossa  ,a fourth     
vacuity is known as the temporal fossa4 
,when the skull of such a mammal as a   
,rabbit1 or ,sheep1 is seen from above1 
there is a vacuity behind the orbits for
 the eyes1 which in life is occupied by 
the muscles which work the lower jaw4   

,it is made by the malar bone extending 
from the back of the orbit and the      
process of bone1 called the zygomatic   
process1 extending forward from the     
articulation of the jaw1 which arches   
out to meet the malar bone4  ,in birds  
there is no conspicuous temporal fossa1 
because the malar bar is a slender rod  
of bone in a line with the lower end of 
the quadrate bone4  ,reptile skulls have
 sometimes one temporal vacuity on each 
side1 as among tortoises1 formed by a   
single lateral bar4 ,these vacuities1   
which correspond to those of mammals in 
position1 are seen from the top of the  
head1 as lateral vacuities behind the   
orbits of the eyes1 and are termed      
superior temporal vacuities4 ,in        
addition to these there is often in     
other reptiles a lateral opening behind 
the eye1 termed the inferior temporal   
vacuity1 seen in ,crocodiles1 in        
,hatteria1 and in ,lizards2 and in such 
skulls there are two temporal bars seen 
in side view1 distinguished as superior 

and inferior4 ,the superior arch always 
includes the squamosal bone1 which is at
 the back of the single bar in mammals4 
,the lower arch includes the malar bone1
 which is in front in the single arch of
 mammals4 ,the circumstance that both   
these arches are connected with the     
quadrate bone makes the double temporal 
arch eminently reptilian4  ,in          
,ornithosaurs the lateral temporal      
vacuity varies from a typically         
reptilian condition to one which1       
without becoming avian1 approaches the  
bird type4 ,in skulls from the ,lias1   
,dimorphodon and ,campylognathus1 there 
is a close parallel to the living ,new  
,zealand reptile ,hatteria1 in the      
vertical position of the quadrate bone  
and in the large size of the vacuity    
behind and below the eye1 which extends 
nearly the height of the skull4 ,in the 
species of the genus ,pterodactylus1 the
 forward inclination of the quadrate    
bone recalls the ,curlew1 ,snipe1 and   
other birds4 ,the back of the head is   

rounded1 and the squamosal bone1 which  
appears to enter into the wall of the   
brain case as in birds and mammals1 is  
produced more outward than in birds1 but
 less than in mammals1 so as to         
contribute a little to the arch which is
 in the position of the post-frontal    
bone of reptiles4 ,it is triangular1 and
 stretches from the outer angle of the  
frontal bone at the back of the orbit to
 the squamosal behind1 where it also    
meets the quadrate bone4 ,its third     
lower branch meets the quadratojugal1   
which rests upon the front of the       
quadrate bone1 as in ,iguanodon1 and is 
unlike ,dimorphodon in its connexions4  
,in that genus the supra-temporal bone1 
or post-orbital bone1 appears to rest   
upon the post-frontal and connect it    
with the quadrato-jugal4 ,in            
,dimorphodon the malar bone is entirely 
removed from the quadrate1 but in       
,pterodactylus it meets its articular   
end4 ,between the post-frontal bone     
above and the quadrato-jugal bone below 

is a small lunate opening1 which        
represents the lateral temporal vacuity2
 and so far1 this is a reptilian        
character4 ,but if the thin post-frontal
 bone were absorbed1 ,pterodactylus     
would resemble birds4 ,there is no      
evidence that the quadrate bone is free 
in any ,ornithosaurs1 as it is in all   
birds1 while in ,dimorphodon it unites  
by suture with the squamosal bone4 ,in  
,ornithostoma the lateral temporal      
vacuity is little more than a slit      
between the quadrate bone below1 the    
quadrato-jugal in front1 and what may be
 the post-frontal bone behind 7see ,fig4
 #b1 p4 #ab74   ,,bones ,,about ,,the   
,,brain  ,the bones containing the brain
 appear to be the same as form the brain
 case in birds4 ,the form of the back of
 the skull varies in two ways4 ,first it
 may be flat above and flat at the back1
 when the back of the head appears to be
 square4 ,this condition is seen in all 
the long-tailed genera1 such as         
,campylognathus from the ,lias and      

,rhamphorhynchus1 and is associated with
 a high position for the upper temporal 
bar4 ,secondly1 the back of the head may
 be rounded convexly1 both above and    
behind4 ,that condition is seen in the  
short-tailed genera1 such as            
,pterodactylus4 ,but in the large       
,cretaceous types1 such as              
,ornithocheirus and ,ornithostoma1 the  
superior longitudinal ridge which runs  
back in the middle line of the face     
becomes elevated and compressed from    
side to side at the back of the head as 
a narrow deep crest1 prolonged backward 
over the neck vertebrae for some inches 
of length4 ,all these three types are   
paralleled more or less in birds which  
have the back of the head square like   
the ,heron1 or rounded like the         
,woodpecker2 or crested1 though the     
crest of the ,cormorant is not quite    
identical with ,ornithocheirus1 being a 
distinct bone at the back of the head in
 the bird which never blends with the   
skull4 ,in so far as the crest is       

reptilian it suggests the remarkable    
crest of the ,chameleon4 ,in the        
structure of the back of the skull the  
bones are a modification of the         
reptilian type of ,hatteria in the ,lias
 genus ,campylognathus1 but the         
reptilian characters appear to be lost  
in the less perfectly preserved skulls  
of ,cretaceous genera4  ,the palate is  
well known in the chief groups of       
,ornithosaurs1 such as ,campylognathus1 
,scaphognathus1 and ,cycnorhamphus4     
,mr4 ,e4 ,t4 ,newton1 ,f4,r4,s41 has    
shown that in the ,english skull from   
the ,lias of ,whitby1 the forms of the  
bones are similar to the palate in birds
 and unlike the conditions in reptiles4 
,there is one feature1 however1 which   
may indicate a resemblance to ,dicynodon
 and other fossil reptiles from ,south  
,africa4 ,a slender bone extends from   
the base of the brain case1 named the   
basi-sphenoid bone1 outward and forward 
to the inner margin of the quadrate bone
 7,fig4 #bb74 ,a bone is found thus     

placed in those ,south ,african         
,reptiles1 which show many resemblances 
to the ,monotreme and ,marsupial        
,mammals4 ,it is not an ordinary element
 of the skeleton and is unknown in      
living animals of any kind in that      
position4 ,it has been thought possible 
that it may represent one of the bones  
which among mammals are diminutive and  
are included in the internal ear4 ,the  
resemblance may have some interest      
hereafter1 as helping to show that      
certain affinities of the ,ornithosaurs 
may lie outside the groups of existing  
reptiles4 ,instead of being directed    
transversely outward1 as in the palatal 
region of ,dicynodon lacerticeps1 they  
diverge outward and forward to the inner
 border of the articulation for the     
lower jaw which is upon the quadrate    
bone4    ,illustration3 ,,fig4 #bb      
,,bones ,,of ,,the ,,palate  ,there is a
 pair of bones which extend forward from
 these inner articular borders of the   
quadrate bones1 and converge in a long  

,v-shape till they merge in the hard    
palate formed by the bones of the front 
of the beak1 named intermaxillary and   
maxillary bones4 ,the limits of the     
bones of the palate are not distinct1   
but there can be no doubt that the front
 of the ,v is the bone named vomer1 that
 the palatine bones are at its sides1   
and that its hinder parts are the       
pterygoid bones as in birds4 ,there is a
 long1 wide1 four-sided1 open space in  
the middle of the palate1 between the   
vomer and the basi-sphenoid bone1 unlike
 anything in birds or other animals4    
,professor ,marsh1 in a figure of the   
palate in the great skull of the        
toothless ,pterodactyle named           
,ornithostoma 7,pteranodon71 from the   
,chalk of ,kansas1 found a large oval   
vacuity in this region of the palate4   
,in that genus the pterygoid bones meet 
each other between the quadrate bones as
 in ,dicynodon 7,fig4 #gc1 p4 #ahb74    
,hence the great palatal vacuity here   
seen in the ,ornithosaur is paralleled  

by the small vacuity in the ,south      
,african reptile1 which is sometimes    
distinct and sometimes partly separated 
from the anterior part of the vacuity   
which forms the openings of the nostrils
 on the palate4  ,the ,solenhofen skulls
 which give any evidence of the palate  
are exposed in side view only1 and the  
bones1 imperfectly seen through the     
lateral vacuities1 are displaced by     
crushing4 ,they include long strips like
 the vomerine bones in the ,lias fossil1
 and they diverge in the same way as    
they extend back to the quadrate bones4 
,the oblique division into vomer in     
front and pterygoid bone behind is shown
 by ,goldfuss in his original figure of 
,scaphognathus4 ,thus there is some     
reason for believing that all           
,ornithosaurs have the palate formed    
upon the same general plan1 which is on 
the whole peculiar to the group1        
especially in not having the palatal    
openings of the nares divided in the    
middle line4 ,it would appear probable  

that the short-tailed animals have the  
pterygoid bones meeting in the middle   
line and triangular2 and that they are  
slender rods entirely separate from each
 other in the long-tailed genera4       
,,the ,,teeth  ,the teeth are all of    
pointed1 elongated shape1 without       
distinction into the kinds seen in most 
mammals and named incisors1 canines1 and
 grinders4 ,they are organs for         
grasping1 like the teeth of the         
fish-eating ,crocodile of ,india1 and   
are not unlike the simple teeth of some 
,porpoises4 ,they are often implanted in
 oblique oval sockets with raised       
borders1 usually at some distance apart 
from each other1 and have the crown     
pointed1 flattened more on the outer    
side than on the inner side1 usually    
directed forward and curved inward4 ,as 
in many extinct animals allied to       
existing reptiles1 the teeth are        
reproduced by germs1 which originate on 
the inner side of the root and grow till
 they gradually absorb the substance of 

the old tooth1 forming a new one in its 
place4 ,frequently in ,solenhofen       
genera1 like ,scaphognathus and         
,pterodactylus1 the successional tooth  
is seen in the jaw on the hinder border 
of the tooth in use4 ,there is some     
variation in the character of bluntness 
or sharpness of the crowns in the       
different genera1 and in their size4    
,the name ,dimorphodon1 given to the    
animal from the ,lias of ,lyme ,regis1  
expresses the fact that the teeth are of
 two kinds4 ,in the front of the jaw    
three or four large long teeth are found
 in the intermaxillary bone on each     
side1 as in some ,plesiosaurs1 while the
 teeth found further back in the        
maxillary bone are smaller1 and directed
 more vertically downward4 ,this        
difference is more marked in the lower  
jaw than in the upper jaw4 ,in          
,rhamphorhynchus the teeth are all      
relatively long and large1 and directed 
obliquely forward1 but absent from the  
extremities of the beak1 as in the      

,german genus from the ,lias named      
,dorygnathus1 in which the bone of the  
lower jaw 7which alone is known7        
terminates in a compressed spear4 ,in   
,scaphognathus the teeth are few1 more  
vertical1 and do not extend backward so 
far as in ,rhamphorhynchus1 but are     
carried forward to the extremity of the 
blunt1 deep jaw4  ,in the short-tailed  
,pterodactyles the teeth are smaller1   
shorter1 wider at the base of the crown1
 closer together1 and do not extend so  
far backward in the jaw4 ,in            
,ornithocheirus two teeth always project
 forward from the front of the jaw4     
,ornithostoma is toothless4   ,,supposed
 ,,horny ,,beak  ,sometimes a horny     
covering has been suggested for the     
beak1 like that seen in birds or        
turtles1 but no such structure has been 
preserved1 even in the ,solenhofen      
,slate1 in which such a structure would 
seem as likely to be preserved as a wing
 membrane1 though there is one doubtful 
exception4 ,there are marks of fine     

blood vessels on some of the jaws1      
indicating a tough covering to the bone4
 ,in ,rhamphorhynchus the jaws appear to
 gape towards their extremities as      
though the interspace had originally    
been occupied by organic substance like 
a horny beak4   ,,lower ,,jaw  ,the     
lower jaw varies in relative length with
 the vertical or horizontal position of 
the quadrate bone in the skull4 ,in     
,dimorphodon the jaw is as long as the  
skull2 but in the genera from the       
,oolitic rocks the mandible is somewhat 
shorter1 and in ,ornithostoma the       
discrepancy reaches its maximum4 ,the   
hinder part of the jaw is never         
prolonged backward much beyond the      
articulation1 differing in this respect 
from ,crocodiles and ,plesiosaurs4  ,the
 depth of the jaw varies4 ,it is slender
 in ,pterodactylus1 and is probably     
stronger relatively to the skull in     
,scaphognathus than in any other form4  
,it fits between the teeth and bones of 
the alveolar border in the skull1 in all

 the genera4 ,in ,dimorphodon its hinder
 border is partly covered by the        
descending edge of the malar process    
which these animals develop in common   
with some ,dinosaurs1 and some          
,anomodont reptiles1 and many of the    
lower mammals4 ,in this hinder region   
the lower jaw is sometimes perforated1  
in the same way as in ,crocodiles4 ,that
 condition is observed in ,dimorphodon1 
but is not found in ,pterodactylus4 ,the
 lower jaw is always composite1 being   
formed by several bones1 as among       
reptiles and birds4 ,the teeth are in   
the dentary bone or bones1 and these    
bones are almost always blended as in   
most birds and ,turtles1 and not        
separate from each other as among       
,crocodiles1 ,lizards1 and ,serpents4   
,an interesting contour for the lower   
border of the jaw is seen in            
,ornithostoma1 as made known in figures 
of ,american examples by ,professors    
,marsh and ,williston4 ,it deepens as it
 extends backwards for two-thirds its   

length1 stops at an angle1 and then the 
depth diminishes to the articulation    
with the skull4 ,this angle of the lower
 jaw is a characteristic feature of the 
jaws of ,mammals4 ,it is seen in the    
monotreme ,echidna1 and is              
characteristic of some ,theriodont      
,reptiles from ,south ,africa1 which in 
many ways resemble ,mammals4 ,the       
character is not seen in the jaws of    
specimens from the ,oolitic rocks1 but  
is developed in the toothed             
,ornithocheirus from the ,cambridge     
,greensand1 and is absent from the jaws 
of existing reptiles and birds4         
,illustration3 ,,fig4 #bc  ,,comparison 
,,of ,,the ,,lower ,,jaw ,,in ,,echidna 
,,and   ,,ornithostoma   ,,summary ,,of 
,,characters ,,of ,,the ,,head  ,taken  
as a whole1 the head differs from other 
types of animals in a blending of       
characters which at the present day are 
found among ,birds and ,reptiles1 with  
some structures which occur in extinct  
groups of animals with similar          

affinities1 and perhaps a slight        
indication of features common to the    
lowest mammals4 ,it is chiefly upon the 
head that the diverse views of earlier  
writers have been based4 ,cuvier was    
impressed with the reptilian aspect of  
the teeth2 but in later times           
discoveries were made of ,birds with    
teeth--,archaeopteryx1 ,ichthyornis1    
,hesperornis4 ,the teeth are quite      
reptilian1 being not unlike miniature   
teeth of ,mosasaurus4 ,if those birds   
had been found prior to the discovery of
 ,pterodactyles1 the teeth might have   
been regarded as a link with the more   
ancient birds1 rather than a crucial    
difference between birds and reptiles4  
,all the specimens show a lateral       
temporal hole in the bones behind the   
eye1 and this is found in no bird or    
mammal1 and is typical of such reptiles 
as ,hatteria4 ,the quadrate bone may not
 be so decisive as ,cuvier thought it to
 be1 for its form is not unlike the     
quadrate of a bird1 and different1 so   

far as ,i have seen1 from that of living
 reptiles4 ,this region of the head is  
reptilian1 and if it occurred in a bird 
the character would be as astonishing as
 was the discovery of teeth in extinct  
birds4 ,these characters of the head are
 also found in fossil animals named     
,dinosaurs1 in association with many    
resemblances to birds in their bones4   
,the palate might conceivably be derived
 from that of ,hatteria by enlarging the
 small opening in the middle line in    
that reptile till it extended forward   
between the vomera2 but it is more      
easily compared with a bird1 which the  
animal resembles in its beak1 and in the
 position of the nares4 ,excepting      
certain ,lizards1 all true existing     
,reptiles have the nostrils far forward 
and bordered by two premaxillary bones  
instead of one intermaxillary1 as in    
,birds and ,ornithosaurs4 ,if nothing   
were known of the animal but its head   
bones1 it would be placed between       
,reptiles and ,birds4     ,,chapter ,,ix

  ,,the ,,backbone1 ,,or ,,vertebral    
,,column   ,the backbone is a more      
deep-seated part of the skeleton than   
the head4 ,it is more protected by its  
position1 and has less varied functions 
to perform4 ,therefore it varies less in
 distinctive character within the limits
 of each of the classes of vertebrate   
animals than either the head or limbs4  
,it is divided into neck bones1 the     
cervical vertebrae2 back bones1 the     
dorsal vertebrae2 loin bones1 the lumbar
 vertebrae2 the sacrum1 or sacral       
vertebrae1 which support the hind limbs2
 and the tail4 ,of these parts the tail 
is the least important1 though it       
reaches a length in existing reptiles   
which sometimes exceeds the whole of the
 remainder of the body1 and includes    
hundreds of vertebrae4 ,it attains its  
maximum among serpents and lizards4 ,in 
frogs it is practically absent4 ,in some
 of the higher mammals it is a rudiment1
 which does not extend beyond the soft  
parts of the body4   ,,the ,,neck  ,the 

neck is more liable to vary than the    
back1 with the habit of life of the     
animal4 ,and although mammals almost    
always preserve the same number of seven
 bones in the neck1 the bones vary in   
length between the short condition of   
the porpoise1 in which the neck is      
almost lost1 and the long bones which   
form the neck of the ,llama1 though even
 these may be exceeded by some fossil   
reptiles like ,tanystrophoeus4 ,in many 
mammals the neck bones do not differ in 
length or size from those of the back4  
,in others1 like the ,horse and ,ox1    
they are much broader and larger4       
,there is the same sort of variation in 
the bones of the neck among birds1 some 
being slender like the ,heron1 others   
broad like the ,swan4 ,but there is also
 a singular variation in number of      
vertebral bones in a bird's neck4 ,at   
fewest there are nine1 which equals the 
exceptionally large number found among  
mammals in the neck of one of the       
,sloths4 ,usually birds have ten to     

fifteen cervical vertebrae1 and in the  
,swan there are twenty-three4 ,most of  
the neck bones of birds are relatively  
long1 and the length of the neck is     
often greater than the remainder of the 
vertebral column4  ,reptiles usually    
have short necks4 ,the common ,turtle   
has eight bones in the neck1 ten in the 
back4 ,the two regions are sharply      
defined by the dorsal shield4 ,their    
articular ends are sometimes cupped in  
front1 in the neck1 sometimes cupped    
behind1 or convex at both ends1 or even 
flattened1 or the articulation may be   
made exceptionally by the neural arch   
alone4 ,nine is the largest number of   
neck bones in existing ,lizards1 and    
there are usually nine in ,crocodiles2  
so that reptiles closely approach       
mammals in number of the neck bones4 ,it
 is remarkable that the maximum number  
in a mammal and in living reptiles      
should coincide with the minimum number 
in birds4 ,therefore the number of      
cervical vertebrae as an attribute of   

,mammal1 ,bird1 or ,reptile1 can only be
 important from its constancy4  ,german 
naturalists affirm on clear evidence    
that the ,solenhofen ,pterodactyles have
 seven cervical vertebrae4 ,in many     
specimens there can be no doubt about   
the number1 because the neck bones are  
easily distinguished from those of the  
back by their size2 but the number is   
not always easy to count4  ,as in       
,birds1 the first vertebra1 or atlas1 in
 ,pterodactyles is extremely short1 and 
is generally--if not always--blended    
with the much longer second vertebra1   
named the axis4 ,the front of the atlas 
forms a small rounded cup to articulate 
with the rounded ball of the            
basioccipital bone at the back of the   
skull4 ,the third and fourth vertebrae  
are longer1 but the length visibly      
shortens in the sixth and seventh4      
,sometimes the vertebrae are slender and
 devoid of strong spinous processes4    
,this is the condition in the little    
,pterodactylus longirostris and in the  

comparatively large ,cycnorhamphus      
,fraasii1 in which there is a slight    
median ridge along the upper surface of 
the arch of the vertebra4 ,this         
condition is paralleled in birds with   
long necks1 especially wading birds such
 as the ,heron4 ,other ,ornithosaurs1   
such as ,ornithocheirus from the        
,cretaceous rocks1 have the neck much   
more massive4 ,the vertebrae are        
flattened on the under side4 ,the arch  
above the nervous matter of the spinal  
cord has a more or less considerable    
transverse expansion1 and may even be as
 wide as long4 ,these vertebrae have    
proportions and form such as may be seen
 in ,vultures or in the ,swan4 ,in      
either case the form of the neck bones  
is more or less bird-like1 and the      
neural spine may be elevated1 especially
 in ,pterodactyles with long tails4     
,one of the most distinctive features of
 the neck bones of a bird is the way in 
which the cervical ribs are blended with
 the vertebrae4 ,they are small1 and    

each is often prolonged in a needle-like
 rod at the side of the neck bone4  ,in 
,ornithocheirus the cervical rib        
similarly blends with the vertebra by   
two articulations1 as in mammals1 so    
that it might escape notice but for the 
channel of a blood vessel which is thus 
inclosed4 ,in several of the older      
,pterodactyles from ,solenhofen the ribs
 of the neck vertebrae remain separated1
 as in a ,crocodile1 though still       
bird-like in their form1 anterior       
position1 and mode of attachment4 ,in   
,terrapins and ,tortoises the long neck 
vertebrae have no cervical ribs4        
,illustration3 ,,fig4 #bd  ,,united     
,,atlas ,,and ,,axis ,,of               
,,ornithocheirus   7,cambridge          
,greensand7  ,the articular surfaces    
between the bodies of the vertebrae1 in 
the neck1 are transversely oval4 ,the   
middle part of this articular joint is  
made by the body of the vertebra2 its   
outer parts are in the neural arch4 ,in 
front this surface is a hollow channel1 

often more depressed than in any other  
animals4 ,the corresponding surface     
behind is convex1 with a process on each
 side at its lower outer angles 7,fig4  
#be74 ,it is a modification of the      
cup-and-ball form of vertebral          
articulation1 which at the present day  
is eminently reptilian4 ,serpents and   
,crocodiles have the articulations      
similarly vertical1 but in both the form
 of the articulation is a circle4 ,in   
,lizards the articular cup is usually   
rather wider than deep1 when the cup and
 ball are developed in the vertebrae2 it
 differs from the vertical condition in 
pterodactyles in being oblique and much 
narrower from side to side4 ,only among 
,crocodiles and ,hatteria is there a    
double articulation for the cervical    
rib1 though in neither order have rib or
 vertebra in the neck the bird-like     
proportions which are usual in these    
animals4 ,pterodactyles show no         
resemblance to birds in this vertebral  
articulation4 ,a ,bird has the          

corresponding surface concave from side 
to side in front1 but it is also convex 
from above downward1 producing what is  
known as the saddle-shaped form which is
 peculiarly avian1 being found in       
existing birds except in part of the    
back in ,penguins4 ,it is faintly       
approximated to in one or two neck      
vertebrae in man4 ,professor ,williston 
remarks that in the toothless           
,pterodactyles of ,kansas the hinder    
ball of the vertebral articulation is   
continued downward and outward as a     
concave articulation upon the processes 
at its outer corners4 ,there are no     
mammals with a cup-and-ball articulation
 between the vertebrae1 so that for what
 it is worth the character now described
 in ,ornithosaurs is reptilian1 when    
judged by comparison with existing      
animals4    ,illustration3 ,,fig4 #be4  
,,cervical ,,vertebra ,,of              
,,ornithocheirus   ,from the ,cambridge 
,greensand  ,low down on each side of   
the vertebra1 at the junction of its    

body with the neural arch1 is a large   
ovate foramen1 transversely elongated1  
and often a little impressed at the     
border1 which is the entrance of the air
 cell into the bone4 ,these foramina are
 often one-third of the length of the   
neck vertebrae in specimens from the    
,cambridge ,greensand1 where the neck   
bones vary from three-quarters of an    
inch to about two and a half inches in  
length1 and in extreme forms are as wide
 as long4 ,the width of the interspace  
between the foramina is one-half the    
width of the vertebrae1 though this     
character varies with different genera  
and species4 ,several species from the  
,solenhofen ,slate have the neck long   
and slender1 on the type of the         
,flamingo4 ,in others the neck is thick 
and short--in the ,scaphognathus        
crassirostris and ,pterodactylus        
spectabilis4 ,some genera with slender  
necks have the bones preserved with a   
curved contour1 such as might suggest a 
neck carried like that of a ,llama or a 

,camel4 ,the neck is occasionally       
preserved in a curve like a capital ,s1 
as though about to be darted forward    
like that of a bird in the act of       
striking its prey4 ,the genera of       
,pterodactyles with short necks may have
 had as great mobility of neck as is    
found among birds named ,ducks and      
,divers2 but those ,pterodactyles with  
stout necks1 such as ,dimorphodon and   
,ornithocheirus1 in which the vertebrae 
are large1 appear to have been built    
more for strength than activity1 and the
 neck bones have been chiefly concerned 
in the muscular effort to use the       
fighting power of the jaws in the best  
way4   ,,the ,,back  ,the region of the 
back in a ,pterodactyle is short as     
compared with the neck1 and relatively  
is never longer than the corresponding  
region in a bird4 ,the shortness results
 partly from the short length of the    
vertebrae1 each of which is about as    
long as wide4 ,there is also a moderate 
number of bones in the back4 ,in most   

skeletons from ,solenhofen these        
vertebrae between the neck and girdle of
 hip bones number from twelve to        
sixteen4 ,they have a general           
resemblance in form to the dorsal       
vertebrae in birds4 ,the greatest number
 of such vertebrae in birds is eleven4  
,the number is small because some of the
 later vertebrae in birds are overlapped
 by the bones of the hip girdle1 which  
extend forward and cover them at the    
sides1 so that they become blended with 
the sacrum4 ,this region of the skeleton
 in the ,dimorphodon from the ,lias is  
remarkable for the length of the median 
process1 named the neural spine1 which  
is prolonged upward like the spines of  
the early dorsal vertebrae of ,horses1  
,deer1 and other mammals4 ,in this      
character they differ from living       
reptiles1 and parallel some ,dinosaurs  
from the ,weald4 ,the bones of the back 
in ,ornithocheirus from the ,cambridge  
,greensand show the under side to be    
well rounded1 so that the articular     

surfaces between the vertebrae1 though  
still rather wider than deep1 are much  
less depressed than in the region of the
 neck4 ,the neural canal for the spinal 
cord has become larger and higher1 and  
the sides of the bone are somewhat      
compressed4 ,strong transverse processes
 for the support of the ribs are        
elevated above the level of the neural  
canal1 at the sides of vertebrae        
compressed on the under sides1 and      
directed outward4 ,between these lateral
 horizontal platforms is the compressed 
median neural spine1 which varies in    
vertical height4 ,the articulation of   
the ribs is not seen clearly4 ,isolated 
ribs from the ,stonesfield ,slate have  
double-headed dorsal ribs1 like those of
 birds4 ,in some specimens from the     
,solenhofen ,slate like the             
,scaphognathus1 in the ,university      
,museum at ,bonn1 dorsal ribs appear to 
be attached by a notch in the transverse
 process of the dorsal vertebra1 which  
resembles the condition in ,crocodiles4 

,variations in the mode of attachment of
 ribs among mammals may show that       
character to be of subordinate          
importance4 ,von ,meyer has described   
the first pair of ribs as frequently    
larger than the others1 and there appear
 in ,rhamphorhynchus to be examples     
preserved of the sternal ribs1 which    
connect the dorsal ribs with the        
sternum4 ,six pairs have been counted4  
,a more interesting feature in the ribs 
consists in the presence behind the     
sternum1 which is shorter than the      
corresponding bone in most birds1 of    
median sternal ribs4 ,they are slender  
,v-shaped bones in the middle line of   
the abdomen1 which overlapped the ends  
of the dorsal ribs like the similar     
sternal bones of reptiles4 ,such        
structures are unknown among ,birds and 
,mammals4 ,there is no trace in the     
dorsal ribs of the claw-like process1   
which extends laterally from rib to rib 
as a marked feature in many birds4 ,its 
presence or absence may not be          

important1 because it is represented by 
fibro-cartilage in the ribs of          
crocodiles1 and may be a small cartilage
 near the head of the rib in serpents1  
and is only ossified in some ribs of the
 ,new ,zealand reptile ,hatteria4 ,so   
that it might have been present in a    
fossil animal without being ossified and
 preserved4 ,although the structure is  
associated with birds1 it is possibly   
also represented by the great bony      
plates which cover the ribs in          
,chelonians1 and combine to form the    
shield which covers the turtle's back4  
,the structure is as characteristic of  
reptiles as of birds1 but is not        
necessarily associated with either4     
,illustration3 ,,fig4 #bf    ,the upper 
figures show the side and back of a     
dorsal vertebra of   ,ornithocheirus    
compared with corresponding views of the
 side and   back of a dorsal vertebra of
 a ,crocodile  ,there are two remarkable
 modifications of the early dorsal      
vertebrae in some of the ,cretaceous    

,pterodactyles4 ,first1 in the genus    
,ornithodesmus from the ,weald the early
 dorsal vertebrae are blended together  
into a continuous mass1 like that which 
is found in the corresponding region of 
the living ,frigate-bird1 only more     
consolidated1 and similar to that       
consolidated structure found behind the 
dorsal vertebrae1 known as the sacrum1  
made by the blending of the vertebrae   
into a solid mass which supports the hip
 bones4 ,secondly1 in some of the       
,cretaceous genera of ,pterodactyles of 
,europe and ,america the vertebrae in   
the front part of the back are similarly
 blended1 but their union is less       
complete2 and in genera ,ornithocheirus 
and ,ornithostoma--the former chiefly   
,english1 the latter chiefly            
,american--the sides of the neural      
spines are flattened to form an oval    
articular surface on each side1 which   
gives attachment to the flattened ends  
of their shoulder-blade bones named the 
scapulae4 ,this condition is found in no

 other animals4 ,three vertebrae appear 
to have their neural arches thus united 
together4 ,the structure so formed may  
be named the notarium to distinguish it 
from the sacrum4   ,,sacrum  ,for some  
mysterious reason the part of the       
backbone which lies between the bones of
 the hips and supports them is termed   
the sacrum4 ,among living reptiles the  
number of vertebrae in this region is   
usually two1 as in lizards and          
crocodiles4 ,there are other groups of  
fossil reptiles in which the number of  
sacral vertebrae is in some cases less  
and in other cases more4 ,there is1     
perhaps1 no group in which the sacrum   
makes a nearer approach to that of birds
 than is found among these              
,pterodactyles1 although there are more 
sacral vertebrae in some ,dinosaurs4 ,in
 birds the sacral vertebrae number from 
five to twenty-two4 ,in bats the number 
is from five to six4 ,in some           
,solenhofen species1 such as            
,pterodactylus dubius and ,p4 ,kochi and

 ,p4 grandipelvis1 the number is usually
 five or six4 ,the vertebrae are        
completely blended4 ,the pneumatic      
foramina in the sacrum1 so far as they  
have been observed1 are on the under    
sides of the transverse processes2 while
 in the corresponding notarial structure
 in the shoulder girdle the foramina are
 in front of the transverse processes4  
,almost any placental mammal in which   
the vertebrae of the sacral region are  
anchylosed together has a similar       
sacrum1 which differs from that of birds
 in the more complete individuality of  
the constituent bones remaining evident4
 ,the transverse processes in front of  
the sacrum are wider than in its hinder 
part2 so that the pelvic bones which are
 attached to it converge as they extend 
backward1 as among mammals4 ,the bodies 
of the vertebrae forming the sacrum are 
similar in length to those of the back4 
,each transverse process is given off   
opposite the body of its own vertebra1  
but from a lower lateral position than  

in the region of the back1 in which the 
vertebrae are free4    ,illustration3   
,,fig4 #bg4  ,,sacrum ,,of              
,,rhamphorhynchus    ,showing the       
complete blending of the vertebrae and  
ribs as in a bird1   with the           
well-defined ,iliac bones1 produced     
chiefly in front of the   acetabulum for
 the head of the femur4  ,the hip bones 
are closely united with the sacrum by   
bony union1 and rarely appear to come   
away from the sacral vertebrae1 as among
 mammals and reptiles1 though this      
happens with the ,lias ,pterodactyles4  
,in the ,stonesfield ,slate and         
,solenhofen ,slate the slender          
transverse processes from the vertebrae 
blend with the ilium of the hip girdle1 
and form a series of transverse foramina
 on each side of the bodies of the      
vertebrae4 ,in the ,cambridge ,greensand
 genera the part of the ilium above the 
acetabulum for the articular head of the
 femur appears to be always broken away1
 so that the relation of the sacrum to  

the pelvis has not been observed4 ,this 
character is no mark of affinity1 but   
only shows that ossification obliterated
 sutures among these animals in the same
 way as among birds4  ,the great        
difference between the sacrum of a      
,pterodactyle and that of a bird has    
been rendered intelligible by the       
excellent discussion of the sacral      
region in birds made by ,professor      
,huxley4 ,he showed that it is only the 
middle part of the sacrum of a chicken  
which corresponds to the true sacrum of 
a reptile1 and comprises the five       
shortest of the vertebrae2 while the    
four in front correspond to those of the
 lower part of the back1 which either   
bear no ribs or very short ribs1 and are
 known as the lumbar region in mammals1 
so that the lower part of the back      
becomes blended with the sacrum1 and    
thus reduces the number of dorsal       
vertebrae4 ,similarly the five vertebrae
 which follow the true sacral vertebrae 
are originally part of the tail1 and    

have been blended with the other        
vertebrae in front1 in consequence of   
the extension along them of the bird's  
hip bones4 ,this interpretation helps to
 account for the great length of the    
sacrum in many birds1 and also explains 
in part the singular shortness of the   
tail in existing birds4 ,the            
,ornithosaur sacrum has neither the     
lumbar nor the caudal portions of the   
sacrum of a bird4    ,,the ,,tail  ,the 
tail is perhaps the least important part
 of the skeleton1 since it varies in    
character and length in different       
genera4 ,the short tails seen in typical
 pterodactyles include as few as ten    
vertebrae in ,pterodactylus grandipelvis
 and ,p4 ,kochi1 and as many as fifteen 
vertebrae in ,pterodactylus             
longirostris4 ,the tails are more like  
those of mammals than existing birds1 in
 which there are usually from six to ten
 vertebrae terminating in the           
ploughshare bone4 ,but just as some     
fossil birds1 like the ,archaeopteryx1  

have about twenty long and slender      
vertebrae in the tail1 so in the        
pterodactyle ,rhamphorhynchus this      
region becomes greatly extended1 and    
includes from thirty-eight to forty     
vertebrae4 ,in ,dimorphodon the tail    
vertebrae are slightly fewer4 ,the      
earliest are very short1 and then they  
become elongated to two or three times  
the length of the early tail vertebrae1 
and finally shorten again towards the   
extremity of the tail1 where the bones  
are very slender4 ,in all long-tailed   
,ornithosaurians the vertebrae are      
supported and bordered by slender       
ossified ligaments1 which extend like   
threads down the tail1 just as they do  
in ,rats and many other mammals and in  
some lizards4  ,professor ,marsh was    
able to show that the extremity of the  
tail in ,rhamphorhynchus sometimes      
expands into a strong terminal caudal   
membrane of four-sided somewhat         
rhomboidal shape4 ,he regards this      
membrane as having been placed          

vertically4 ,it is supported by delicate
 processes which represent the neural   
spines of the vertebrae prolonged       
upward4 ,they are about fifteen in      
number4 ,a corresponding series of      
spines on the lower border1 termed      
chevron bones1 equally long1 were given 
off from the junctions of the vertebrae 
on their under sides1 and produced      
downward4 ,this vertical appendage is of
 some interest because its expansion is 
like the tail of a fish4 ,it suggests   
the possibility of having been used in a
 similar way to the caudal fin as an    
organ for locomotion in water1 though it
 is possible that it may have also      
formed an organ used in flight for      
steering in the air4    ,illustration3  
,,fig4 #bh4  ,,extremity ,,of ,,the     
,,tail ,,of ,,rhamphorhynchus           
,,phyllurus 7,,marsh7    ,showing the   
processes on the upper and under sides  
of the vertebrae   which make the       
terminal leaf-like expansion  ,the tail 
vertebrae from the ,cambridge ,greensand

 are mostly found isolated or with not  
more than four joints in association4   
,they are very like the slender type of 
neck vertebrae seen in long-necked      
pterodactyles1 but are depressed1 and   
though somewhat wider are not unlike the
 tail vertebrae of the ,rhamphorhynchus4
 ,the pneumatic foramen in them is a    
mere puncture4 ,they have no transverse 
processes or neural spines1 nor         
indications of ribs1 or chevron bones4  
,the hindermost specimens of tail       
vertebrae observed have the neural arch 
preserved to the end1 as among reptiles2
 whereas in mammals this arch becomes   
lost towards the end of the tail4 ,the  
processes by which the vertebrae are    
yoked together are small4 ,there is     
nothing to suggest that the tail was    
long1 except the circumstance that the  
slender caudal vertebrae are almost as  
long as the stout cervical vertebrae in 
the same animal4 ,no small caudal       
vertebrae have ever been found in the   
,cambridge ,greensand4 ,the tail is very

 short1 according to ,professor         
,williston1 in the toothless            
,ornithostoma in the ,chalk of ,kansas4 
,,chapter ,x  ,,the ,,hip-,,girdle ,,and
 ,,hind ,,limb   ,the bones of the      
hip-girdle form a basin which incloses  
and protects the abdominal vital organs4
 ,it consists on each side of a         
composite bone1 the unnamed bones--ossa 
innominata of the older                 
anatomists--which are each attached to  
the sacrum on their inner side1 and on  
the outer side give attachment to the   
hind limbs4 ,as a rule three bones enter
 into the borders of this cup1 termed   
the acetabulum1 in which the head of the
 thigh bone1 named the ,femur1 moves    
with a more or less rotary motion4      
,there are a few exceptions in this     
division of the cup between three bones1
 chiefly among ,salamanders and certain 
,frogs4 ,in ,crocodiles the bone below  
the acetabular cup is not divided into  
two parts4 ,and in certain ,plesiosaurs 
from the ,oxford                        

,clay--,muraenosaurus--the actual       
articulation appears to be made by two  
bones--the ilium and ischium4 ,the three
 bones which form each side of the      
pelvis are known as the ilium1 or hip   
bone1 sometimes termed the aitch bone2  
secondly1 the ischium1 or sitz bone1    
being the bone by which the body is     
supported in a sitting position2 and    
thirdly the pubis1 which is the bone in 
front of the acetabulum4 ,the pubic     
bones meet in the middle line of the    
body on the under side of the pelvis in 
man1 and on each side are partly        
separated from the ischia by a foramen1 
spoken of as the obturator foramen1     
which in ,pterodactyles is minute and   
almost invisible1 when it exists4       
,there is often a fourth bony element in
 the pelvis4 ,in some ,salamanders a    
single cartilage is directed forward1   
and forked in front4 ,according to      
,professor ,huxley something of this    
kind is seen in the ,dog4 ,the pair of  
bones which extend forward in front of  

the pelvis in ,crocodiles may be of the 
same kind1 in which case they should be 
called prepubic bones4 ,but among the   
lower mammals named marsupials a pouch  
is developed for the protection of the  
young and supported by two slender bones
 attached to the pubes1 and these bones 
have long been known as marsupial bones4
 ,in a still lower group of mammalia    
named monotremata1 which lay eggs1 and  
in many ways approximate to reptiles and
 birds1 stronger bones are developed on 
the front edge of the pubes1 and termed 
prepubic bones4 ,they do not support a  
marsupium4  ,naturalists have been      
uncertain as to the number of bones in  
the pelvis of ,pterodactyles1 because   
the bones blend together early in life1 
as in birds4 ,some follow the ,amphibian
 nomenclature1 and unite the ischium and
 pubis into one bone1 which is then     
termed ischium1 when the prepubis is    
termed the pubis1 and regarded as       
removed from the acetabulum4 ,there is  
no ground for this interpretation1 for  

the sutures are clear between the three 
pelvic bones in the acetabulum in some  
specimens1 like ,cycnorhamphus ,fraasii1
 from ,solenhofen1 and some examples of 
,ornithocheirus from the ,cambridge     
,greensand4 ,pterodactyles all have     
prepubic bones1 which are only known in 
,ornithorhynchus and ,echidna among     
mammals1 and are absent from the higher 
mammals and birds4 ,they are unknown in 
any other existing animals1 unless      
present in ,crocodiles1 in which ischium
 and pubis are always undivided4        
,therefore it is interesting to examine 
the characters of the ,ornithosaurian   
pelvis4  ,the acetabulum for the head of
 the femur is imperforate1 being a      
simple oval basin1 as in ,chelonian     
reptiles and the higher ,mammals4 ,it   
never shows the mark of the ligamentous 
attachment to the head of the femur1    
which is seen in ,mammals4 ,in ,birds   
the acetabulum is perforated1 as in many
 of the fossils named ,dinosaurs1 and in
 ,monotremata4    ,illustration3 ,,fig4 

#bi4  ,,comparison ,,of ,,the ,,left    
,,side ,,of ,,the ,,pelvis ,,in   ,a    
,,bird ,,and ,a ,,pterodactyle          
,secondly1 the ilium is elongated1 and  
extends quite as much in front of the   
acetabulum as behind it4 ,the bone is   
not very deep in this front process4    
,among existing animals this relation of
 the bone is nearer to birds than to any
 other type1 since birds alone have the 
ilium extended from the acetabulum in   
both directions4 ,the form of the       
,pterodactyle ilium is usually that of  
the embryo bird1 and its slender        
processes compare in relative length    
better with those of the unhatched fowl 
and ,apteryx of ,new ,zealand than with 
the plate-like form in adult birds4  ,in
 mammals the ilium is directed forward1 
and even in the ,cape ant-eater         
,orycteropus there is only an           
inappreciable production of the bone    
backward behind the acetabulum4 ,among  
reptiles the general position of the    
acetabulum is at the forward termination

 of the ilium1 though the ,crocodile has
 some extension of the bone in both     
directions1 without forming distinct    
anterior and posterior processes4 ,this 
anterior and posterior extension of the 
ilium is seen in the ,theriodont        
reptiles of ,russia and of ,south       
,africa1 as well as in ,dinosaurs4      
,illustration3 ,,fig4 #cj4  ,,left      
,,pelvic ,,bones ,,with ,,prepubic      
,,bone ,,in   ,,pterodactylus           
,,longirostris  ,thirdly1 in all        
pterodactyles the ischium and pubis are 
more or less completely blended into a  
sheet of bone1 unbroken by perforation1 
though there is usually a minute        
vascular foramen2 or the lower border   
may be notched between the ischium and  
the pubis1 as in some of the ,solenhofen
 species1 and the pubis does not reach  
the median line of the body4 ,but in    
,dimorphodon the pelvic sheet of bone is
 unbroken by any notch or perforation4  
,the notch between the ischium and pubis
 is well marked in ,pterodactylus       

longirostris1 and better marked in      
,pterodactylus dubius1 ,cycnorhamphus   
,fraasii1 and ,rhamphorhynchus4 ,the    
fossil animals which appear to come     
nearest to the ,pterodactyles in the    
structure of the pelvis are ,theriodonts
 from the ,permian rocks of ,russia4    
,the type known as ,rhopalodon has the  
ilium less prolonged front and back1 and
 is much deeper than in any             
,pterodactyle2 but the acetabulum is    
imperforate1 and the ischium and pubis  
are not always completely separated from
 each other by suture4 ,in the pelvis   
referred to the ,theriodont             
,deuterosaurus there is some            
approximation to the pelvis of          
,rhamphorhynchus and of ,pterodactylus  
dubius in the depth of the division     
between the pubis and ischium4          
,illustration3 ,,fig4 #ca  ,,pelvis     
,,and ,,prepubic ,,bones ,,of           
,,rhamphorhynchus    ,on the left-hand  
side the two prepubic bones are         
separate4 ,on the   right-hand they are 

united into a transverse bar which      
overlaps the   front of pelvis seen from
 the under side  ,there are three       
modifications of the ,ornithosaurian    
pelvis4 ,first1 the type of             
,rhamphorhynchus1 in which the pubis and
 ischium are inclined somewhat backward1
 and in which the two prepubic bones are
 triangular1 and are often united       
together to form a transverse bow in    
front of the pubic region4  ,secondly1  
there is the ordinary form of pelvis in 
which the pubis and ischium usually     
unite with each other down their length1
 as in ,dimorphodon1 but sometimes1 as  
in ,pterodactylus dubius1 divide        
immediately below the acetabulum4 ,all  
these types possess the paddle-shaped   
prepubic bones1 which are never united  
in the median line4  ,thirdly1 there is 
the cretaceous form indicated by        
,ornithocheirus and ,ornithostoma1 in   
which the posterior half of the ilium is
 modified in a singular way1 since it is
 more elevated towards the sacrum than  

the anterior half1 suggesting the       
contour of the upper border of the ilium
 in a lizard4 ,without being            
reptilian--the anterior prolongation of 
the bone makes that impossible--it      
suggests the lizards4 ,this type also   
possesses prepubic bones4 ,they appear1 
according to ,professor ,williston1 to  
be more like the paddle-shaped bones of 
,pterodactylus than like the angular    
bones in ,rhamphorhynchus4 ,the prepubic
 bones are united in the median line as 
in ,rhamphorhynchus4 ,but their median  
union in that genus favours the         
conclusion that the bones were united in
 the median line in all species1 though 
they are only co-ossified in these two  
families4    ,illustration3 ,,fig4 #cb4 
,,the ,,pelvic ,,bones ,,of ,,an        
,,alligator ,,seen ,,from   ,,below     
,the bones in front are here regarded as
 prepubic1 but are commonly   named     
pubic  ,this median union of the        
prepubic bones is a difference from     
those mammals like the ,ornithorhynchus 

and ,echidna1 which approach nearest to 
the ,reptilia4 ,in them the prepubic    
bones have a long attachment to the     
front margin of the pubis1 and extend   
their points forward without any        
tendency for the anterior extremities to
 approximate or unite4 ,the marsupial   
mammals have the same character1 keeping
 the marsupial bones completely distinct
 from each other at their free          
extremities4 ,the only existing animals 
in which an approximation is found to   
the prepubic bones in ,pterodactyles are
 ,crocodiles1 in bones which most       
writers term the pubic bones4 ,this     
resemblance1 without showing any strong 
affinity with the ,crocodilia1 indicates
 that ,crocodiles have more in common   
with the fossil flying animals than any 
other group of existing reptiles2 for   
other reptiles all want prepubic bones1 
or bones in front of the pubic region4  
,,the ,,hind ,,limb  ,the hind limb is  
exceptionally long in proportion to the 
back4 ,this is conspicuous in the       

skeletons of the short-tailed           
,pterodactyles1 and is also seen in     
,dimorphodon4 ,in ,rhamphorhynchus the  
hind limb is relatively much shorter1 so
 that the animal1 when on all fours1 may
 have had an appearance not unlike a    
,bat in similar position4 ,the limb is  
exceptionally short in the little       
,ptenodracon brevirostris4 ,the bones of
 the hind limb are exceptionally        
interesting4 ,one remarkable feature    
common to all the specimens is the great
 elongation of the shin bones relatively
 to the thigh bones4 ,the femur is      
sometimes little more than half the     
length of the tibia1 and always shorter 
than that bone4 ,the proportions are    
those of mammals and birds4 ,some       
mammals have the leg shorter than the   
thigh1 but mammals and birds alone1     
among existing animals1 have the        
proportions which characterise          
,pterodactyles4 ,the foot appears to    
have been applied to the ground not     
always as in a bird1 but more often in  

the manner of reptiles1 or mammals in   
which the digits terminate in claws4    
,,the ,,femur    ,illustration3 ,,fig4  
#cc4  ,,the ,,femur    ,on the right is 
a front view of femur of a bear4 ,in the
 middle are   front and side views of   
the femur of ,ornithocheirus4 ,on the   
left is   the femur of ,echidna4 ,these 
comparisons illustrate the mammalian    
characters of the ,pterodactyle thigh   
bone  ,the thigh bone1 on account of the
 small size of many of the specimens1 is
 not always quite clear evidence as an  
indication of technical resemblance to  
other animals4 ,the bone is always a    
little curved1 has always a rounded1    
articular head1 and rounded distal      
condyles4 ,its most remarkable features 
are shown in the large1 well-preserved  
specimens from the ,cambridge           
,greensand4 ,the rounded1 articular head
 is associated with a constricted neck  
to the bone1 followed by a comparatively
 straight shaft with distal condyles1   
less thickened than in mammals4 ,no bird

 is known1 much less any reptile1 with a
 femur like ,ornithocheirus4 ,only among
 ,mammals is a similar bone known with a
 distinct neck2 and only a few mammals  
have the exceptional characters of the  
rounded head and constricted neck at all
 similar to the ,cretaceous             
,pterodactyles4 ,a few types1 such as   
the higher apes1 the ,hyrax1 and animals
 especially active in the hind limb1    
have a femur at all resembling the      
,pterodactyle in the pit for the        
obturator externus muscle1 behind the   
trochanter major1 such as is seen in a  
small femur from ,ashwell4 ,the femur   
varies in different genera1 so as to    
suggest a number of mammalia rather than
 any particular animal for comparison4  
,these approximations may be            
consequences of the ways in which the   
bones are used4 ,when functional        
modifications of the skeleton are       
developed1 so as to produce similar     
forms of bones1 the muscles to which    
they give attachment1 which act upon the

 bones1 and determine their growth1 are 
substantially the same4 ,in the         
,pterodactylus longirostris the femur   
corresponds in length to about eleven   
dorsal vertebrae4 ,the end next the shin
 bone is less expanded than is usual    
among ,mammals1 and rather suggests an  
approach to the condition in            
,crocodiles1 in the moderate thickness  
and breadth of the articular end1 and   
the slight development of the terminal  
pulley-joint4 ,one striking feature of  
the femur is the circumstance that the  
articular head1 as compared with the    
distal end1 is directed forward and very
 slightly inward and upward4 ,so that   
allowing for the outward divergence of  
the pelvic bones1 as they extend        
forward1 there must have been a tendency
 to a knock-kneed approximation of the  
lower ends of the thigh bones1 as in    
,mammals and ,birds1 rather than the    
outward divergence seen in ,reptiles4   
,apparently the swing of the leg and    
foot1 as it hung on the distal end of   

the femur1 must have tended rather to an
 inward than to an outward direction1 so
 that the feet might be put down upon   
the same straight line2 this arrangement
 suggests rapid movement4               
,illustration3 ,,fig4 #cd4  ,,comparison
 ,,of ,,the ,,tibia ,,and ,,fibula ,,in 
,,ornithosaur ,,and ,,vulture   ,,tibia 
,,and ,,fibula  ,in ,pterodactylus      
longirostris the tibia is slender1 more 
than a fifth longer than the femur4 ,a  
crest is never developed at the proximal
 end1 like that seen in the ,guillemot  
and ,diver and other water birds4 ,the  
bone is of comparatively uniform        
thickness down the shaft in most of the 
,solenhofen specimens1 as in most birds4
 ,at the distal end the shin bone       
commonly has a rounded1 articular       
termination1 like that seen in birds4   
,this is conspicuous in the             
,pterodactylus grandis4 ,in other       
specimens the tarsal bones1 which form  
this pulley1 remain distinct from the   
tibia2 and the upper row of these bones 

appears to consist of two bones1 like   
those which in many ,dinosaurs combine  
to form the pulley-like end of the tibia
 which represents the bird's drum-stick 
bone4 ,they correspond with the ankle   
bones in man named astragalus and os    
calcis4  ,complete ,english specimens of
 tibia and fibula are found in the genus
 ,dimorphodon from the ,lias1 in which  
the terminal pulley of the distal end   
has some expansion1 and is placed       
forward towards the front of the tibia1 
as in some birds4 ,the rounded surface  
of the pulley is rather better marked   
than in birds4 ,the proximal end of the 
shaft is relatively stout1 and is       
modified by the well-developed fibula1  
which is a short external splint bone   
limited to the upper half of the tibia1 
as in birds2 but contributing with it to
 form the articular surface for the     
support of the lower end of the femur1  
taking a larger share in that work than 
in birds4 ,frequently there is no trace 
of the fibula visible in ,solenhofen    

specimens as preserved2 or it is        
extremely slender and bird-like1 as in  
,pterodactylus longirostris4 ,in        
,rhamphorhynchus it appears to extend   
the entire length of the tibia1 as in   
,dinosaurs4 ,in the specimens from the  
,cambridge ,greensand there is          
indication of a small proximal crest to 
the tibia with a slight ridge1 but no   
evidence that this is due to a separate 
ossification4 ,the patella1 or knee-cap1
 is not recognised in any fossil of the 
group4 ,there is no indication of a     
fibula in the specimens thus far known  
from the ,chalk rocks either of ,kansas 
in ,america1 or in ,england4  ,the      
region of the tarsus varies from the    
circumstance that in many specimens the 
tibia terminates downward in a rounded  
pulley1 like the drum-stick of a bird2  
while in other specimens this union of  
the proximal row of the tarsal bones    
with the tibia does not take place1 and 
then there are two rows of separate     
tarsal bones1 usually with two bones in 

each row4 ,when the upper row is united 
with the tibia the lower row remains    
distinct from the metatarsus1 though no 
one has examined these separate tarsal  
bones so as to define them4   ,,the     
,,foot    ,illustration3 ,,fig4 #ce4    
,,metatarsus ,,and ,,digits ,,in ,,three
 ,,types ,,of   ,,ornithosaurs  ,the    
foot sometimes has four toes1 and       
sometimes five4 ,there are four somewhat
 elongated1 slender metatarsal bones1   
which are separate from each other and  
never blended together1 as in birds4    
,there has been a suspicion that the    
metatarsal bones were separate in the   
young ,archaeopteryx4 ,in the young of  
many birds the row of tarsal bones at   
the proximal end of the metatarsus comes
 away1 and there is a partial division  
between the metatarsal bones1 though    
they remain united in the middle4 ,and  
among ,penguins1 in which the foot bones
 are applied to the ground instead of   
being carried in the erect position of  
ordinary birds1 there is always a       

partial separation between the          
metatarsal bones1 though they become    
blended together4 ,the ,pterodactyle is 
therefore different from birds in       
preserving the bones distinct through   
life1 and this character is more like   
,reptiles than ,mammals4 ,the individual
 bones are not like those of ,dinosaurs1
 and diverge in ,rhamphorhynchus as     
though the animals were web-footed4     
,there is commonly a rudimentary fifth  
metatarsal4 ,it is sometimes only a     
claw-shaped appendage1 like that seen in
 the ,crocodile4 ,it is sometimes a     
short bone1 completely formed1 and      
carrying two phalanges in ,solenhofen   
specimens3 though no trace of these     
phalanges is seen in the large toothless
 ,pterodactyles from the ,cretaceous    
rocks of ,north ,america4 ,in the       
,pterodactylus longirostris the number  
of foot bones on the ordinary digits is 
two1 three1 four1 five1 as in lizards2  
but the short fifth metatarsal has only 
two toe bones4 ,in ,dimorphodon the     

fifth digit was bent upward1 and        
supported a membrane for flight4 ,there 
are slight variations in the number of  
foot bones4 ,in the species             
,pterodactylus scolopaciceps the number 
of bones in the toes follows the formula
 two1 three1 three1 four4 ,in           
,pterodactylus micronyx the number is   
two1 three1 three1 three4 ,the terminal 
claws are much less developed than is   
usual with ,birds2 and there is a       
difference from ,bats in the unequal    
length of the digits4 ,taken as a whole1
 the foot is perhaps more reptilian than
 avian1 and in some genera is           
crocodilian4  ,the foot is the light    
foot of an active animal4 ,von ,meyer   
thought that the hind legs were too     
slender to enable the animal to walk on 
land2 and ,professor ,williston1 of the 
,university of ,kansas1 remarks that the
 rudimentary claws and weak toes        
indicate that the animal could not have 
used the feet effectively for grasping1 
while the exceedingly free movement of  

the femur indicates great freedom of    
movement of the hind legs2 and he       
concludes that the function of the legs 
was chiefly for guidance in flight      
through their control over the          
movements1 and expresses his belief that
 the animal could not have stood upon   
the ground with its feet4 ,there may be 
evidence to sustain other views4 ,if the
 limb bones are reconstructed1 they form
 limbs not wanting in elegance or       
length4 ,if it is true1 as ,professor   
,williston suggests1 that the weight of 
his largest animals with the head three 
feet long1 and a stretch of wing of     
eighteen or nineteen feet1 did not      
exceed twenty pounds1 there can be no   
objection to regarding these animals as 
quadrupeds1 or even as bipeds1 on the   
ground of the limbs lacking the strength
 necessary to support the body4 ,the    
slender toes of many birds1 and even the
 two toes of the ostrich1 may be thought
 to give less adequate support for those
 animals than the metatarsals and digits

 of ,pterodactyles4     ,,chapter ,,xi  
,,shoulder-,,girdle ,,and ,,fore ,,limb 
,,sternum  ,the sternum is always a     
distinguishing part of the bony         
structure of the breast4 ,in ,crocodiles
 it is a cartilage to which the sternal 
ribs unite2 and upon its front portion a
 flat knife-like bone called the        
interclavicle is placed4 ,in lizards    
like the ,chameleon1 it is a            
lozenge-shaped structure of thin bony   
texture1 also bearing a long            
interclavicle1 which supports the       
clavicular bones1 named collar bones in 
man1 which extend outward to the        
shoulder blades4 ,among mammals the     
sternum is usually narrow and flat1 and 
often consists of many successive pieces
 in the middle line1 on the under side  
of the body4 ,among ,bats the anterior  
part is somewhat widened from side to   
side1 to give attachment to the collar  
bones1 but the sternum still remains a  
narrow bone1 much narrower than in      
,dolphins1 and not differing in         

character from many other ,mammals1     
notwithstanding the ,bat's power of     
flight4 ,the bone develops a median keel
 for the attachment of the muscles of   
the breast1 but something similar is    
seen in burrowing ,insectivorous mammals
 like the ,moles4 ,so that1 as ,von     
,meyer remarked1 the presence of a keel 
on the sternum is not in itself         
sufficient evidence to prove flight4    
,among birds the sternum is greatly     
developed4 ,broad and short in the      
,ostrich tribe1 it is devoid of a keel2 
and therefore the keel1 if present in a 
bird1 is suggestive of flight4 ,the keel
 is differently developed according to  
the mode of attachment of the several   
pectoral muscles which cover a bird's   
breast4 ,in several water birds the keel
 is strongly developed in front1 and    
dies away towards the hinder part of the
 sternum1 as in the ,cormorant and its  
allies4 ,the sternum in ,german         
,pterodactyles is most nearly comparable
 to these birds4    ,illustration3      

,,fig4 #cf4  ,,comparison ,,of ,,the    
,,sternum  ,in the ,solenhofen ,slate   
the sternum is fairly well preserved in 
many ,ornithosaurs4 ,it is relatively   
shorter than in birds1 and is broader   
than long2 but not very like the sternum
 of reptile or mammal in form4 ,the keel
 is limited to the anterior part of the 
shield of the sternum1 as in ,merganser 
and the ,cormorant1 and is prolonged    
forward for some distance in advance of 
it4 ,von ,meyer noticed the resemblance 
of this anterior process to the         
interclavicle of the ,crocodile in      
position2 but it is more like the keel  
of a bird's sternum1 and is not a       
separate bone as in ,reptiles4 ,in      
,pterodactyles from the ,cretaceous     
rocks1 the side bones1 called coracoids1
 are articulated to saddle-shaped       
surfaces at the hinder part of the base 
of this keel1 which are parallel in     
,ornithocheirus1 as in most birds1 but  
overlap in ,ornithodesmus1 as in ,herons
 and wading birds4    ,illustration3    

,,fig4 #cg4  ,,sternum ,,in             
,,ornithocheirus ,,from ,,the           
,,cambridge   ,,greensand    ,showing   
the strong keel and the facets for the  
coracoid bones on its   hinder border   
above the lateral constrictions  ,the   
keel was pneumatic1 and when broken is  
seen to be hollow1 and appears to have  
been exceptionally high in              
,rhamphorhynchus1 a genus in which the  
wing bones are greatly elongated4 ,von  
,meyer found in ,rhamphorhynchus on each
 side of the sternum a separate lateral 
plate with six pairs of sternal ribs1   
which unite the sternum with the dorsal 
ribs1 as in the young of some birds4    
,the hinder surface of the sternum is   
imperfectly preserved in the toothless  
,pterodactyles of ,kansas4 ,professor   
,williston states that the bone is      
extremely thin and pentagonal in        
outline1 projecting in front of the     
coracoids1 in a stout1 blunt1 keel-like 
process1 similar to that seen in the    
,pterodactyles of the ,cambridge        

,greensand4 ,american specimens have not
 the same notch behind the articulation 
for the coracoid to separate it from the
 transverse lateral expansion of the    
sternal shield4 ,the lateral margin in  
the ,cambridge ,greensand specimens     
figured by ,professor ,owen and myself  
is broken2 but ,professor ,williston had
 the good fortune to find on the margin 
of the sternum the articular surfaces   
which gave attachment to the sternal    
ribs4 ,the margin of the sternal bone   
thickens at these facets1 four of which 
are preserved4 ,the sternum in          
,ornithostoma was about four and a half 
inches long by less than five and a half
 inches wide4 ,the median keel extends  
forward for rather less than two inches1
 while in the smaller ,cambridge species
 of ,ornithocheirus it extends forward  
for less than an inch and a half4  ,a   
sternum of this kind is unlike that of  
any other animal1 but has most in common
 with a bird2 and may be regarded as    
indicating considerable power of flight4

 ,the bone cannot be entirely attributed
 to the effect of flight1 since there is
 no such expanded sternal shield in     
,bats4 ,the small number of sternal ribs
 is even more characteristic of birds   
than mammals or reptiles4   ,,the       
,,shoulder-,,girdle  ,the bones which   
support the fore limb are one of the    
distinctive regions of the skeleton     
defining the animal's place in nature4  
,among most of the lower vertebrata1    
such as ,amphibians and ,reptiles1 the  
girdle is a double arch--the arch of the
 collar bone or clavicles in front1 and 
the arch of the shoulder-blade or       
scapula behind4 ,the clavicular arch1   
when it exists1 is formed of three or   
five parts--a medium bar named the      
interclavicle1 external to which is a   
pair of bones called clavicles1 reaching
 to the front of the scapulae when they 
are present2 and occasionally there is a
 second pair of bones called            
supraclavicles1 extending from the      
clavicles up the front margins of the   

scapulae4 ,thus the clavicular arch is  
placed in front of the scapular arch4   
,the supraclavicles are absent from all 
living ,reptiles1 and the clavicles are 
absent from ,crocodiles4 ,the           
interclavicle is absent from all mammals
 except ,echidna and ,ornithorhynchus4  
,clavicles also may be absent in some   
orders of mammals4 ,hence the clavicular
 arch may be lost1 though the collar    
bones are retained in man4  ,the        
scapular arch also is more complicated  
and more important in the lower than in 
the higher vertebrata4 ,it may include  
three bones on each side named coracoid1
 precoracoid1 and scapula4 ,but in most 
vertebrates the coracoid and precoracoid
 appear never to have been segmented so 
as to be separated from each other2 and 
it is only among extinct types of       
reptiles1 which appear to approximate to
 the ,monotreme mammals1 that separate  
precoracoid bones are found2 though     
among most mammals1 probably1 there are 
stages of early development in which    

precoracoids are represented by small   
cartilages1 though few mammals except   
,edentata like the ,sloths and          
,ant-eaters1 retain even the coracoids  
as distinct bones4 ,therefore1 excepting
 the ,edentata and the ,monotremes1 the 
distinctive feature of the mammalian    
shoulder-girdle appears to be that the  
limbs are supported by the              
shoulder-blades1 termed the scapulae4   
,among reptiles there are several       
distinct types of shoulder-girdle4      
,chelonians possess a pair of bones     
termed coracoids which have no connexion
 with a sternum2 and their scapulae are 
formed of two widely divergent bars1    
divided by a deeper notch than is found 
in any fossil reptiles4 ,among ,lizards 
both scapula and coracoid are widely    
expanded1 and the coracoid is always    
attached to the sternum4 ,chameleons    
have the blade of the scapula long and  
slender1 but the coracoid is always as  
broad as it is long4 ,crocodiles have   
the bone more elongated1 so that it has 

somewhat the aspect of a very strong    
first sternal rib when seen on the      
ventral face of the animal4 ,the bone is
 perforated by a foramen1 which would   
probably lie in the line of separation  
from the precoracoid if any such        
separation had ever taken place4 ,the   
scapula1 or shoulder-blade1 of          
,crocodiles is a similar flat bone1 very
 much shorter than the scapula of a     
,chameleon1 and more like that of the   
,new ,zealand ,hatteria4 ,thus there is 
very little in common between the       
several reptilian types of              
shoulder-girdle4    ,illustration3      
,,fig4 #ch4  ,,comparison ,,of ,,scapula
 ,,and ,,coracoid ,,in ,,three          
,,pterodactyles ,,and ,a ,,bird  ,in    
birds the apparatus for the support of  
the wings has a far-off resemblance to  
the ,crocodilian type4 ,the coracoid    
bones1 instead of being directed        
laterally outward and upward from the   
sternum1 as among ,crocodiles1 are      
directed forward1 so as to prolong the  

line of the breast bone1 named the      
sternum4 ,the bird's coracoid is        
sometimes flattened towards the breast  
bone among ,swans and other birds2 yet  
as a rule the coracoid is a slender bar1
 which combines with the still more     
slender and delicate blade of the       
scapula1 which rests on the ribs1 to    
make the articulation for the upper arm 
bone4 ,among reptiles the scapula and   
coracoid are more or less in the same   
straight line1 as in the ,ostrich1 but  
in birds of flight they meet at an angle
 which is less than a right angle1 and  
where they come in contact the external 
surface is thickened and excavated to   
make the articulation for the head of   
the humerus4 ,there is nothing like this
 shoulder-girdle outside the class of   
birds1 until it is compared with the    
corresponding structure in these extinct
 animals called ,pterodactyles4 ,the    
resemblance between the two is          
surprising4 ,it is not merely the       
identity of form in the coracoid bone   

and the scapula1 but the similar angle  
at which they meet and the similar      
position of the articulation for the    
humerus4 ,everything in the             
,pterodactyle's shoulder-girdle is      
bird-like1 except the absence of the    
representative of the clavicles1 that   
forked ,v-shaped bone of the bird which 
in scientific language is known as the  
furculum1 and is popularly termed the   
merry-thought4 ,this kind of            
shoulder-girdle is found in the genera  
from the ,lias and the ,oolitic rocks1  
both of this country and ,germany4  ,in 
the ,cretaceous rocks the scapula       
presents1 in most cases1 a different    
appearance4 ,the coracoid is an         
elongated1 somewhat triangular bone1    
compressed on the outer margin as in    
birds1 but differing alike from birds   
and other ,pterodactyles in not being   
prolonged forward beyond the            
articulation for the humerus4 ,in these 
,cretaceous genera1 toothed and         
toothless alike1 the articulation for   

the upper arm bone truncates the        
extremity of the coracoid1 so that the  
bone is less like that of a bird in this
 feature4 ,perhaps it shows a           
modification towards the crocodilian    
direction4 ,the scapula1 which unites   
with the coracoid at about a right      
angle1 is similarly truncated by the    
articular surface for the humerus2 but  
the bone is somewhat expanded           
immediately beyond the articulation1 and
 compressed2 and instead of being       
directed backward1 it is directed inward
 over the ribs to articulate with the   
neural arches of the early dorsal       
vertebrae in the genera found in strata 
associated with the ,chalk4 ,as the bone
 approaches this articulation1 it       
thickens and widens a little1 becoming  
suddenly truncated by an ovate facet1   
which exactly corresponds to the        
transversely ovate impression1 concave  
from front to back1 which is seen in the
 neural arches of the dorsal vertebrae  
on which it fits4 ,this condition is not

 present in all ,cretaceous             
,pterodactyles4 ,it does not occur in   
the ,kansas fossil1 named by ,professor 
,marsh1 ,nyctodactylus4 ,and it appears 
to be absent from the ,pterodactyles of 
the ,english ,weald1 named              
,ornithodesmus4    ,illustration3 ,,fig4
 #ci4  ,,the ,,notarium    ,an          
ossification which gives attachment to  
the scapulae seen in the   early dorsal 
vertebra of ,ornithocheirus    7,from   
the ,cambridge ,greensand7              
,illustration3 ,,fig4 #dj4              
,,restoration ,,of ,,the                
,,shoulder-,,girdle ,,in ,,the          
,,cretaceous ,,ornithocheirus           
,showing how the scapulae articulate    
with a vertebra and the   articulation  
of the coracoids with the sternum4 ,the 
humeral   articulation with the coracoid
 is unlike the condition shown   in     
other ,ornithosaurs  ,there is no       
approach to this transverse position of 
the scapulae among birds4 ,and while the
 form of the bones in the older genera  

of ,ornithosaurs is singularly          
bird-like1 the angular arrangement in   
this ,cretaceous genus is obtained by   
closely approximating the articulations 
on the sternum1 so that the coracoids   
extend outward as in reptiles1 instead  
of forward as in birds2 and the         
extremities of the scapulae similarly   
approximate towards each other4 ,this   
rather recalls the relative positions of
 scapula and coracoid among crocodiles4 
,if crocodile and bird had been         
primitive types of animals instead of   
surviving types1 it might almost seem as
 though there had been a cunning and    
harmonious blending of one with the     
other in evolving this form of          
shoulder-girdle4   ,,the ,,fore ,,limb  
,the bones of the fore limb1 generally1 
correspond in length with the similar   
parts of the hind limb4 ,the upper arm  
bone corresponds with the upper leg     
bone1 and the fore-arm bone is as long  
as the fore-leg bone2 then differences  
begin4 ,the bones which correspond to   

the back of the hand in man1 termed the 
metacarpus1 are variable in length in   
,pterodactyles--sometimes very long and 
sometimes short4 ,the wing metacarpal   
bone is always stout1 and the others are
 slender4 ,the extremity of the         
metacarpus was applied to the ground4   
,three small digits of the hand are     
developed from the three small          
metacarpal bones1 and terminate in large
 claws4  ,the great wing finger was bent
 backward1 and only touched the ground  
where it fitted upon the wing metacarpal
 bone4 ,it appears sometimes to have    
been as long as the entire vertebral    
column4  ,owing to the circumstance that
 the joint in the arm in ,pterodactyles 
was not at the wrist as among birds1 but
 between the metacarpus and the         
phalanges1 it follows that the fore limb
 was longer than the hind limb when the 
metacarpus was long2 but the difference 
would not interfere with the movements  
of the animal1 either upon four feet or 
on two feet1 for in bats and birds the  

disproportion in length is greater4     
,,humerus ,,or ,,upper ,,arm ,,bone     
,the first bone in the fore-arm1 the    
humerus1 is remarkable chiefly for the  
compressed crescent form of its upper   
articular end1 which is never rounded   
like the head of the upper arm bone in  
man1 and secondly for the great         
development of the external process of  
bone near that end1 termed the radial   
crest4 ,sir ,richard ,owen compared the 
bone to the humerus of both birds and   
crocodiles1 but in its upper articular  
end the crocodile bone may be said to be
 more like a bird than it is like the   
,pterodactyle4 ,in flying reptiles the  
articular surface next to the           
shoulder-girdle is somewhat             
saddle-shaped1 being concave from side  
to side above and convex vertically1    
while most animals with which it can be 
compared have the articular head of the 
bone convex in both directions4 ,a      
remarkable exception to this general    
rule is found in some fossil animals    

from ,south ,africa1 which1 from        
resemblance to mammals in their teeth1  
have been termed ,theriodonts4 ,they    
sometimes have the head of the bone     
concave from side to side and convex in 
the vertical direction4 ,to this        
condition ,ornithorhynchus makes a      
slight approximation4 ,the singular     
expansion of the structure called the   
radial crest finds no close parallel in 
reptiles1 though ,crocodiles have a     
moderate crest on the humerus in the    
same position2 and in ,theriodonts the  
radial crest extends much further down  
the shaft of the humerus4 ,no bird has a
 radial crest of a similar kind1 though 
it is prolonged some way down the shaft 
in ,archaeopteryx4 ,in ,pterodactyles it
 sometimes terminates outward in a      
smooth1 rounded surface1 which might    
have been articular if any structure    
could have articulated with it4 ,there  
is also a moderate expansion of the bone
 on the ulnar side in some              
,pterodactyles1 so that the proximal end

 often incloses nearly three-fourths of 
an ovate outline4 ,the termination of   
the radial crest is at the opposite end 
of this oval to the wider articular part
 of the head of the bone1 in some       
specimens from the ,cambridge           
,greensand4 ,the radial crest is more   
extended in ,rhamphorhynchus4 ,all      
specimens of the humerus show a twist in
 the length of the bone1 so that the end
 towards the fore-arm1 which is wider   
than the shaft1 makes a right angle with
 the radial crest on the proximal end1  
which is not seen in birds4 ,the shaft  
of the humerus is always stouter than   
that of the femur1 though different     
genera differ in this respect4  ,the    
humerus in genera from rocks associated 
with the ,chalk presents two            
modifications1 chiefly seen in the      
characters of the distal end of the     
bone4 ,one of these is a stout bone with
 a curiously truncated end where it     
joins the two bones of the fore-arm2 and
 the other is more or less remarkable   

for the rounded form of the distal      
condyles4 ,both types show distinct     
articular surfaces4 ,the inner one is   
somewhat oblique and concave1 the outer 
one rounded2 the two being separated by 
a concave channel1 so that the ulna     
makes an oblique articulation with the  
bone as in birds1 and the radius        
articulates by a more or less truncated 
or concave surface4    ,illustration3   
,,fig4 #da4  ,,comparison ,,of ,,the    
,,humerus ,,in ,,pterodactyle ,,and     
,,bird   ,,ulna ,,and ,,radius          
,illustration3 ,,fig4 #db4  ,,comparison
 ,,of ,,the ,,bones ,,of ,,the          
,,fore-,,arm ,,in   ,,bird ,,and        
,,ornithosaur  ,the bones of the        
fore-arm are similar to each other in   
size1 and if there be any difference    
between them the ulna is slightly the   
larger4 ,there is some evidence that in 
,rhamphorhynchus the upper end of the   
ulna was placed behind the radius1      
probably in consequence of the mode of  
attachment of those bones to the        

humerus4 ,the ulna abutted towards the  
inner and lower border1 while the radius
 was towards the upper border1          
consequent upon the twist in the        
humerus4 ,this condition corresponds    
substantially with the arrangement in   
birds1 but differs from birds in the    
relatively more important part taken by 
the radius in making the articulation4  
,the bones are compared in ,dimorphodon 
with the ,golden ,eagle drawn of the    
same size 7,fig4 #db74 ,in birds the    
ulna supports the great feathers of the 
wing1 and this may account for the size 
of the bone4 ,the ulna is best seen at  
its proximal end in the specimens from  
the ,cambridge ,greensand1 where there  
is a terminal olecranon ossification    
forming an oblique articulation1 which  
frequently comes away and is lost4 ,it  
is sometimes well preserved1 and        
indicated by a suture4 ,the examples of 
ulna from the ,lias show a slight       
expansion of the bone at both ends1 and 
at the distal end toward the wrist the  

articulation is well defined1 where the 
bone joins the carpus4 ,the larger      
specimens of the bone are broken4 ,the  
distal articular surface is only        
connected with the proximal end of the  
bone in small specimens3 it always shows
 on the one margin a concavity1 followed
 by a prominent boss1 and an oblique    
articulation beyond the boss4 ,on the   
side towards the radius1 on the lower   
end of the shaft there is an angular    
ridge1 which marks the line along which 
the ulna overlaps the radius4 ,the lower
 end of the radius has a simple1        
slightly convex articulation1 somewhat  
bean-shaped4 ,no rotation of these bones
 on each other was possible as in man4  
,there is a third bone in the fore-arm4 
,this bone1 named the pteroid1 is       
commonly seen in skeletons from         
,solenhofen4 ,it was regarded by ,von   
,meyer as having supported the wing     
membrane in flight4 ,some writers have  
interpreted it as an essential part of  
the ,pterodactyle skeleton1 and ,von    

,meyer thought that it might possibly   
indicate a fifth digit in the hand4 ,the
 only existing structure at all like it 
is seen in the ,south ,african          
insectivorous mammal named              
,chrysochloris capensis1 the golden     
mole1 which also has three bones in the 
fore-arm1 the third bone extending      
half-way up towards the humerus4 ,in    
that animal the third bone appears to be
 behind the others and adjacent to the  
ulna4 ,in the ,german fossils the       
pteroid articulated with a separate     
carpal or metacarpal bone1 placed on the
 side of the arm adjacent to the radius1
 and the radius is always more inward   
than the ulna4 ,if the view suggested by
 ,von ,meyer is adopted1 this bone would
 be a first digit extending outward and 
backward towards the humerus4 ,that view
 was adopted by ,professor ,marsh4 ,it  
involves the interpretation of what has 
been termed the lateral carpal as the   
first metacarpal bone1 which would be as
 short as that of a bird1 but turned in 

the opposite direction backward4 ,the   
first digit would then only carry one   
phalange1 and would not terminate in a  
claw1 but lie in the line of the tendon 
which supports the anterior wing        
membrane of a bird4  ,the third bone in 
the fore-arm of ,chrysochloris does not 
appear to correspond to a digit4 ,the   
bone is on the opposite side of the arm 
to the similar bone of a ,pterodactyle1 
and therefore cannot be the same        
structure in the ,golden ,mole4 ,the    
interpretation which makes the pteroid  
bone the first digit has the merit of   
accounting for the fifth digit of the   
hand4 ,all the structures of the hand   
are consistent with this view4 ,the     
circumstance that the bone is rarely    
found in contact with the radius1 but   
diverging from it1 shows that it plays  
the same part in stretching the membrane
 in advance of the arm1 that the fifth  
digit holds in supporting the larger    
wing membrane behind the arm4           
,according to ,professor ,williston1 the

 ,american toothless ,pterodactyle      
,ornithostoma has but a single phalange 
on the corresponding first toe of the   
hind foot1 and that bone he describes as
 long1 cylindrical1 gently curved1 and  
bluntly pointed4 ,there is some support 
for this interpretation2 but ,i have not
 seen any ,english or ,german           
,pterodactyles with only one phalange in
 the first toe4  ,the wing in           
,pterodactyles would thus be stretched  
between two fingers which are bent      
backward1 the three intermediate digits 
terminating in claws4   ,,the ,,carpus  
,the wrist bones in the reptilia usually
 consist of two rows4 ,in ,crocodiles1  
in the upper row there is a large inner 
and a small outer bone1 behind which is 
a lunate bone1 the remainder of the     
carpus being cartilaginous4 ,only one   
carpal is converted into bone in the    
lower row4 ,it is placed immediately    
under the smaller upper carpal4 ,in     
,chelonians1 the turtle and tortoise    
group1 the characters of the carpus vary

 with the family4 ,in the upper row     
there are usually two short carpals1    
which may be blended1 under the ulna2   
while the two under the radius are      
commonly united4 ,the lower row is made 
up of several small bones4 ,lizards1    
too1 usually have three bones in the    
proximal row and five smaller bones in  
the distal row4  ,the correspondence of 
the distal carpals with the several     
metacarpal bones of the middle hand is a
 well-known feature of the structure of 
the wrist4  ,von ,meyer remarks that the
 carpus is made up of two rows of small 
bones in the ,solenhofen ,pterodactyles2
 while in birds there is one row        
consisting of two bones4 ,the structure 
of the carpus is not distinct in all    
,german specimens2 but in the           
short-tailed ,solenhofen genera the     
bones in the two rows retain their      
individuality4  ,in all the ,cretaceous 
genera the carpal bones of each row are 
blended into a single bone1 so that two 
bones are superimposed1 which may be    

termed the proximal and distal carpals4 
,one specimen shows by an indication of 
sutures the original division of the    
distal carpal into three bones2 and the 
separated constituent bones are very    
rarely met with4 ,two bones of the three
 confluent elements contribute to the   
support of the wing metacarpal1 and the 
third gives an articular attachment to  
the bone which extends laterally at the 
inner side of the carpus1 which ,i now  
think may be the first metacarpal bone  
turned backward towards the humerus4    
,the three component bones meet in the  
circular pneumatic foramen in the middle
 of the under side of the distal carpal4
 ,there is no indication of division of 
the proximal carpal in these genera into
 constituent bones4    ,illustration3   
,,fig4 #dc4  ,,carpus ,,from            
,,ornithocheirus   7,cambridge          
,greensand7  ,this condition is somewhat
 different from birds4 ,in #ahgc ,dr4   
,rosenberg1 of ,dorpat1 showed that     
there is in the bird a proximal carpal  

formed of two elements1 and a distal    
carpal also formed of two elements4     
,therefore the two constituents of the  
distal carpal in the bird which blends  
in the mature animal with the           
metacarpus1 forming the rounded pulley  
joint1 may correspond with two of the   
three bones in the ,cretaceous          
,pterodactyle ,ornithocheirus4  ,the    
width of a proximal carpal rarely       
exceeds two inches1 and that of a distal
 carpal is about an inch and            
three-quarters4 ,two such bones when in 
contact would not measure more than one 
inch in depth4 ,the lower surface shows 
that the wing had some rotary movement  
upon the carpus outward and backward4   
,,metacarpus    ,illustration3 ,,fig4   
#dd4  ,,metacarpus ,,in ,,two           
,,ornithosaurs  ,the metacarpus consists
 of bones which correspond to the back  
of the hand4 ,the first digit of the    
hand in clawed animals has the          
metacarpal bone short1 or shorter than  
the others4 ,among mammals metacarpal   

bones are sometimes greatly elongated2  
and a similar condition is found in     
,pterodactyles1 in which the metacarpal 
bone may be much longer than the        
phalange which is attached to it4 ,two  
metacarpal bones appear to be singularly
 stouter than the others4 ,the first    
bone of the first digit1 if rightly     
determined1 is much shorter than the    
others1 and is1 in fact1 no longer than 
the carpus 7,fig4 #dc74 ,it is a flat   
oblong bone1 attached to the inner side 
of the lower carpal1 and instead of     
being prolonged distally in the same    
direction as the other metacarpal bones1
 is turned round and directed upward1 so
 that its upper edge is flush with the  
base of the radius1 and gives attachment
 to a bone which resembles a terminal   
phalange of the wing finger4 ,according 
to this interpretation it is the first  
and only phalange in the first digit4   
,the bone is often about half as long as
 the fore-arm1 terminates upward in a   
point1 is sometimes curved1 and         

frequently diverges outward from the    
bones of the fore-arm1 as preserved in  
the associated skeleton1 being stretched
 towards the radial crest of the        
humerus4 ,this mode of attachment of the
 supposed first metacarpal1 which is    
true for all ,cretaceous pterodactyles1 
has not been shown to be the same for   
all those from the ,solenhofen ,slate4  
,there is no greater anomaly in this    
metacarpal and phalange on the inner    
side being bent backward1 than there is 
in the wing finger being bent backward  
on the outer side4 ,the three slender   
intervening digits extend forward       
between them1 as though they were       
applied to the ground for walking4  ,the
 bone which is usually known as the wing
 metacarpal is frequently stouter at the
 proximal end towards the carpus than   
towards the phalange4 ,at the carpal end
 it is oblong and truncated1 with a     
short middle process1 which may have    
extended into the pit in the base of the
 carpal bone2 while the distal terminal 

end is rounded exactly like a pulley4   
,there is great difference in the length
 of the metacarpus4 ,in the ,american   
genus ,ornithostoma it is much longer   
than the fore-arm4 ,in ,rhamphorhynchus 
it is remarkably short1 though perhaps  
scarcely so short as in ,dimorphodon or 
in ,scaphognathus4 ,the largest         
,cretaceous examples are about two      
inches wide where they join the carpus4 
,the bone is sometimes a little curved4 
,between the first and fifth or wing    
metacarpal are the three slender        
metacarpal bones which give attachment  
to the clawed digits4 ,they bear much   
the same relation to the wing metacarpal
 that the large metatarsal of a         
,kangaroo has to the slender bones of   
the instep which are parallel to it4    
,the facet for the wing metacarpal on   
the carpus is clearly recognised1 but as
 a rule there is no surface with which  
the small metacarpals can be separately 
articulated4 ,one or two exceptional    
specimens from the ,cambridge ,greensand

 appear to have not only surfaces for   
the wing metacarpal1 but two much       
smaller articular surfaces1 giving      
attachment to smaller metacarpals2 while
 in one case there appears to be only   
one of these additional impressions4 ,it
 is certain that all the animals from   
the ,lias and ,oolites have three clawed
 digits1 but at present ,i have seen no 
evidence that there were three in the   
,cretaceous genera1 though ,professor   
,williston's statements and restoration 
appear to show that the toothless       
,pterodactyles have three4 ,another     
difference from the ,oolitic types1     
according to ,professor ,williston1 is  
in the length of the slender metacarpals
 of the clawed phalanges being about    
one-third that of the wing metacarpal1  
but this is probably due to imperfect   
ossification at the proximal end2 for at
 the distal end the bones all terminated
 on the same level1 showing that the    
four outer digits were applied to the   
ground to support the weight of the     

body4 ,the corresponding bone in the    
,horse and ,oxen is carried erect1 so as
 to be in a vertical line with the bones
 of the fore-arm2 and the same position 
prevails usually1 though not invariably1
 with the corresponding bone in the hind
 limb1 while in many clawed mammals the 
metacarpus and metatarsus are both      
applied upon the ground4 ,in            
,pterodactyles the metatarsal bones are 
preserved in the rock in the same       
straight line with the smaller bones of 
the foot1 or make an angle with the shin
 bone1 leading to the conviction that   
the bones of the foot were applied to   
the ground as in ,man1 and sometimes as 
in the ,dog1 and were thus modified for 
leaping4 ,just as the human metacarpus  
is extended in the same line with the   
bones of the fore-arm1 and the movement 
of jointing occurs where the fingers    
join the metacarpus1 so ,pterodactyles  
also had these bones differently        
modified in the fore and hind limbs for 
the functions of life4 ,the result is to

 lengthen the fore limb as compared with
 the hind limb by introducing into it an
 elevation above the ground which       
corresponds to the length of the        
metacarpus1 always supposing that the   
animal commonly assumed the position of 
a quadruped when upon the earth's       
surface4  ,this position of the         
metacarpus is a remarkable difference   
from ,birds1 because when the bird's    
wing is at rest it is folded into three 
portions4 ,the upper arm bone extends   
backward1 the bones of the fore-arm are 
bent upon it so as to extend forward1   
and then at the wrist the third portion1
 which includes the metacarpus and      
finger bones1 is bent backward4 ,so that
 the metacarpus in the ,pterodactyle    
differs from birds in being in the same 
line as the bones of the fore-arm1      
whereas in birds it is in the same line 
with the digit bones of the hand4 ,it is
 worthy of remark that in ,bats1 which  
are so suggestive of ,pterodactyles in  
some features of the hand1 the          

metacarpals and phalanges are in the    
same straight line2 so that in this     
respect the bat is more like the bird4  
,but ,pterodactyles in the relation of  
these bones to flight are quite unlike  
any other animal1 and have nothing in   
common with the existing animals named  
,reptiles4   ,,the ,,hand  ,from what   
has just been said it follows that the  
construction of the hand is unique4 ,it 
may be contrasted with the foot of a    
bird4 ,the bone which is called1 in the 
language of anatomists1 the             
tarso-metatarsus1 and is usually free   
from feathers and covered with skin1 is 
commonly carried erect in birds1 so that
 the whole body is supported upon it2   
and from it the toes diverge outward4   
,it is formed in birds of three separate
 bones blended together4 ,in the fore   
limb of the ,pterodactyle the metacarpus
 has the same relation to the bones of  
the fore-arm that the metatarsus has to 
the corresponding bones of the leg in a 
bird4 ,but the three metacarpal bones in

 the ,pterodactyle remain distinct from 
each other1 perhaps because the main    
work of that region of the skeleton has 
devolved upon the digit called the wing 
finger1 which is not recognised in the  
bird4 ,in the ,pterodactyles from the   
,solenhofen ,slate there is a           
progressive number of phalanges in the  
three small digits of the hand1 which   
were applied to the ground4 ,this number
 in the great majority of species       
follows the formula of two bones in the 
first1 three bones in second1 and four  
in the third2 so that in the innermost  
of the clawed digits only one bone      
intervenes between the metacarpal and   
the claw4 ,the fingers slightly increase
 in length with increase in number of   
bones which form them4    ,illustration3
 ,,fig4 #de4  ,,claw ,,phalange ,,from  
,,the ,,hand ,,in   ,,ornithocheirus4   
7,half natural size7    ,illustration3  
,,fig4 #df4  ,,metacarpus ,,and ,,digits
 ,,of ,,the ,,hand ,,in ,,birds   ,,with
 ,,claws  ,the terminal claw bones are  

unlike the claws of ,birds or ,reptiles4
 ,they are compressed from side to side1
 and extremely deep and strong1 with    
evidence of powerful attachment for     
ligaments1 so that they rather resemble 
in their form and large size the claws  
of some of the carnivorous fossil       
reptiles1 often grouped as ,dinosauria1 
such as have been termed ,aristosuchus  
and ,megalosaurus4 ,in the hand of the  
,ostrich the first and second digits    
terminate in claws1 while the third is  
without a claw4 ,but these claws of the 
,ostrich and other birds are slender1   
curved1 and rather feeble organs4 ,in   
the ,archaeopteryx1 a fossil bird which 
agrees with the ,pterodactyles in       
retaining the separate condition of the 
metacarpal bones and in having the same 
number of phalanges in two of the       
fingers of the fore limb1 the terminal  
claws are rather more compressed from   
side to side1 and stronger than in the  
,ostrich1 but not nearly so strong as in
 the ,pterodactyle4 ,the ,archaeopteryx 

differs from the ,pterodactyle in having
 no trace of a wing finger4 ,the first  
metacarpal bone is short1 as in all     
birds2 and the first phalange scarcely  
lengthens that segment of the first     
digit of the ,bird's hand to the same   
length as the other metacarpal bones4   
,it therefore was not bent backward like
 the first digit in ,pterodactyles4 ,the
 wing finger1 from which the genius of  
,cuvier selected the scientific         
name--,pterodactyle--for these fossils1 
yields their most distinctive character4
 ,it is a feature which could only be   
partly paralleled in the ,bat1 by making
 changes of structure which would remove
 every support to the wing but the      
outermost digit of that animal's hand4  
,in the ,bat's hand the membrane for    
flight is extended chiefly by four      
diverging metacarpal bones4 ,there are  
only two or three phalanges in each     
digit in its four wing fingers4 ,in     
,pterodactyles the metacarpal bones are1
 as we have seen1 arranged in close     

contact1 and take no part in stretching 
the wing4   ,,the ,,wing ,,finger  ,in  
,birds there is nothing whatever to     
represent the wing finger of the        
,pterodactyle1 for it is an organ       
external to the finger bones of the     
bird1 and contains four phalanges4 ,the 
first phalange is quite different from  
the others4 ,its length is astonishing  
when compared with the small phalanges  
of the clawed fingers4 ,the articular   
surface1 which joins on to the wing     
metacarpal bone1 is a concave           
articulation1 which fits the pulley in  
which that bone ends4 ,the pulley       
articulation admits of an extension     
movement in one direction only4 ,many   
specimens show the wing finger to be    
folded up so as to extend backward4 ,the
 whole finger is preserved in other     
specimens straightened out so as to be  
in line with the metacarpus4 ,this      
condition is well seen in ,professor    
,marsh's specimen of ,rhamphorhynchus1  
which has the wing membrane preserved1  

in which all bones of the fore-arm      
metacarpus and wing finger are extended 
in a continuous curve4 ,the outer       
surface of the end of the first bone of 
the wing finger overlaps the wing       
metacarpal1 so that a maximum of        
strength and resistance is provided in  
the bony structures by which the wing is
 supported4 ,there is1 therefore1 in    
flight only one angular bend in the     
limb1 and that is between the upper arm 
bone and the fore-arm4  ,an immense     
pneumatic foramen is situate in a groove
 on the under side of the upper end of  
the first phalange in ,ornithocheirus1  
but is absent in specimens from the     
,kimeridge clay4 ,this bone is long and 
stout4 ,it terminates at the lower end  
in an obliquely truncated articular     
surface4 ,specimens occur in the        
,cambridge ,greensand which are #b      
inches broad at the upper end and nearly
 #a-a_/b inch wide at the lower end4 ,an
 imperfect bone from the ,chalk is      
#ad-a_/b inches long4 ,the bones are all

 flattened4 ,specimens from the ,chalk  
of ,kansas at ,munich are #bh inches    
long4 ,the second phalange is concave at
 the upper articular end and convex in  
the longer direction at the lower end4  
,the articular points of union between  
the several phalanges form prominences  
on the under side of the finger in      
consequence of the adjacent bones being 
a little widened at their junction4 ,it 
should be mentioned that there is a     
proximal epiphysis or separate bone to  
the first phalange1 adjacent to the     
pulley joint of the metacarpal bone1    
which is like the separate olecranon    
process of the ulna of the fore-arm4 ,it
 sometimes comes away in specimens from 
the ,chalk and ,cambridge ,greensand1   
leaving a large circular pit with a     
depressed narrow border4 ,on the outer  
side of this process is a rounded boss1 
which may possibly have supported the   
bone1 if it were applied to the ground  
with the wing folded up1 like the wing  
of a ,bat directed upward and backward  

at the animal's side4  ,the four bones  
of the wing finger usually decrease     
progressively in length1 so that in     
,rhamphorhynchus1 in which the length of
 the animal's head only slightly exceeds
 #c-a_/b inches1 the first phalange is  
nearly as long1 the second phalange is  
about #c-a_/d inches1 the third #b-c_/d 
inches1 and the fourth a little over #b 
inches4 ,thus the entire length of the  
four phalanges slightly exceeds #aa     
inches1 or rather more than three times 
the length of the head4 ,but the        
fore-arm and metacarpus in this type    
only measure #c inches4 ,therefore the  
entire spread of wings could not have   
been more than #b feet #i inches4  ,the 
largest ,ornithosaur in which accurate  
measurements have been made is the      
toothless ,pterodactyle ,ornithostoma1  
also named ,pteranodon1 from ,north     
,america4 ,in that type the head appears
 to have been about #c or #d feet long1 
and the wing finger exceeded #e feet2   
while the length of the fore-arm and    

metacarpus exceeded #c feet4 ,the width 
of the body would not have been more    
than #a foot4 ,the length of the short  
humerus1 which was about #aa inches1 did
 not add greatly to the stretch of the  
wing2 so that the spread of the wings as
 stretched in flight may be given as    
probably not exceeding #ag or #ah feet4 
,a fine example of the wing bones of    
this animal quite as large has been     
obtained by the 7,british ,museum       
,natural ,history74 ,many years ago1 on 
very fragmentary materials1 ,i estimated
 the wings in the ,english ,cretaceous  
,ornithocheirus as probably having a    
stretch of #bj feet in the largest      
specimens1 basing the calculation partly
 upon the extent of the longest wings in
 existing birds relatively to their     
bones1 and partly upon the size of the  
largest associated bones which were then
 known4     ,,chapter ,,xii  ,,evidences
 ,,of ,,the ,,animal',s ,,habits ,,from 
,,its ,,remains   ,such are the more    
remarkable characters of the bones in a 

type of animal life which was more      
anomalous than any other which peopled  
the earth in the ,secondary ,epoch of   
geological time4 ,its skeleton in       
different parts resembles ,reptiles1    
,birds1 and ,mammals2 with modifications
 and combinations so singular that they 
might have been deemed impossible if    
,nature's power of varying the skeleton 
could be limited4 ,since ,ornithosaurs  
were provided with wings1 we may believe
 the animals to some extent to have     
resembled birds in habit4 ,their modes  
of progression were more varied1 for the
 structures indicate an equal capacity  
for movement on land as a biped1 or as a
 quadruped1 with movement in the air4   
,there is little evidence to support the
 idea that they were usually aquatic    
animals4 ,the majority of birds which   
frequent the water have their bodies    
stored with fat and the bones of their  
extremities filled with marrow4 ,and a  
bird's marrow bones are stouter and     
stronger than those which are filled    

with air4 ,there are few1 if any1 bones 
of ,pterodactyles so thick as to suggest
 the conclusion that they contained     
marrow1 and the bones of the extremities
 appear to have been constructed on the 
lightest type found among terrestrial   
birds4 ,their thinness1 except in a few 
specimens from the ,wealden rocks1 is   
marvellous2 and all the later           
,pterodactyles show the arrangement1 as 
in birds1 by which air from the lungs is
 conveyed to the principal bones4 ,no   
,pterodactyle has shown any trace of the
 web-footed condition seen in birds     
which swim on the water1 unless the     
diverging bones of the hind foot in     
,rhamphorhynchus supports that          
inference4 ,the bones of the hind foot  
are relatively small1 and if it were not
 that a bird stands easily upon one     
foot1 might be considered scarcely      
adequate to support the animal in the   
position which terrestrial birds usually
 occupy4 ,yet1 as compared with the     
length and breadth of the foot in an    

,ostrich1 the toes of an ,ornithosaur   
are seen to be ample for support4 ,these
 facts appear to discourage the idea    
that the animals were equally at home on
 land and water1 and in air4  ,some     
light may be thrown upon the animal's   
habits by the geological circumstances  
under which the remains are found4 ,the 
,pterodactyle named ,dimorphodon1 from  
the ,lias of the south of ,england1 is  
associated with evidences of terrestrial
 land animals1 the best known of which  
is ,scelidosaurus1 an armoured ,dinosaur
 adapted by its limbs for progression on
 land4 ,and the ,pterodactyle           
,campylognathus1 from the ,lias of      
,whitby1 is associated with trunks of   
coniferous trees and remains of         
,insects4 ,so that the occurrence of    
,pterodactyles in a marine stratum is   
not inconsistent with their having been 
transported by streams from off the old 
land surface of the ,lias1 on which     
coniferous trees grew and ,dinosaurs    
lived4  ,similar considerations apply to

 the occurrence of the ,rhamphocephalus 
in the ,stonesfield ,slate of ,england4 
,the deposit is not only formed in      
shallow water1 but contains terrestrial 
,insects1 a variety of land plants1 and 
many ,reptiles and other animals which  
lived upon land4 ,the specimens from the
 ,purbeck beds1 again1 are in strata    
which yield a multitude of the spoils of
 a nearly adjacent land surface2 while  
the numerous remains found in the marine
 ,solenhofen ,slate in ,germany are     
similarly associated with abundant      
evidences of varied types of terrestrial
 life4 ,the evidence grows in force from
 its cumulative character4 ,the ,wealden
 beds1 which yield many terrestrial     
reptiles and so much evidence of        
terrestrial vegetation1 and             
shallow-water conditions of disposition1
 have afforded important ,pterodactyle  
remains from the ,isle of ,wight and    
,sussex4  ,the chief ,english deposit in
 which these fossils are found1 the     
,upper ,greensand1 has afforded         

thousands of bones1 battered and broken 
on a shore1 where they have lain in     
little associated groups of remains1    
often becoming overgrown with small     
marine shells4 ,side by side with them  
are found bones of true terrestrial     
,lizards and ,crocodiles of the type of 
the ,gavial of the ,indian rivers1 many 
terrestrial ,dinosaurs1 and other       
evidences of land life1 including fossil
 resins1 such as are met with in the    
form of amber or copal at the present   
day4  ,the great bones of ,pterodactyles
 found in the ,chalk of ,kent1 near     
,rochester1 became entombed1 beyond     
question1 far from a land surface4      
,there is nothing to show whether the   
animals died on land and were drifted   
out to sea like the timber which is     
found water-logged and sunken after     
being drilled by the ship-worm 7,teredo7
 of that epoch4 ,seeing the power of    
flight which the animal possessed1      
storms may have struck down travellers  
from time to time1 when far from land4  

,evidence of habit of another kind may  
be found in their teeth4 ,they are      
brightly enamelled1 sharp1 formidable2  
and are frequently long1 overlapping the
 sides of the jaws4 ,they are organs    
which are often better adapted for      
grasping than for tearing1 as may be    
seen in the inclined teeth of           
,rhamphocephalus of the ,stonesfield    
,slate2 and better adapted for killing  
than tearing1 from their piercing forms 
and cutting edges1 in genera like       
,ornithocheirus of the ,greensand4 ,the 
manner in which the teeth were implanted
 and carried is better paralleled by the
 fish-eating crocodile of ,indian rivers
 than by the flesh-eating crocodiles1 or
 ,muggers1 which live indifferently in  
rivers and the sea4 ,as the ,kingfisher 
finds its food 7see ,fig4 #bj7 from the 
surface of the water without being in   
the common sense of the term a water    
bird1 so some ,pterodactyles may have   
fed on fish1 for which their teeth are  
well adapted1 both in the stream and by 

the shore4  ,a ,pterodactyle's teeth    
vary a good deal in appearance4 ,the few
 large teeth in the front of the jaw in 
,dimorphodon1 associated with the many  
small vertical teeth placed further     
backward1 suggest that the taking of    
food may have been a process requiring  
leisure1 since the hinder teeth adapted 
to mincing the animal's meat are        
extremely small4 ,the way in which the  
teeth are shaped and arranged differs   
with the genera4 ,in ,pterodactylus they
 are short and broad and few1 placed for
 the most part towards the front of the 
jaws4 ,their lancet-shaped form         
indicates a shear-like action adapted to
 dividing flesh4 ,in the associated     
genus ,rhamphorhynchus the teeth are    
absent from the extremity of the jaw1   
are slender1 pointed1 spaced far apart1 
and extend far backward4 ,when the jaws 
of the ,rhamphorhynchus are brought     
together there is always a gap between  
them in front1 which has led to belief  
that the teeth were replaced by some    

kind of horny armature which has        
perished4 ,in the long-nosed ,english   
type of ,ornithocheirus the jaws are    
compressed together1 so that the teeth  
of the opposite sides are parallel to   
each other1 with the margins well filled
 with teeth1 which are never in close   
contact1 though occasionally closer and 
larger in front1 in some of the forms   
with thick truncated snouts4  ,it is not
 the least interesting circumstance of  
the dentition of ,pterodactyles that1   
associated in the same deposits with    
these most recent genera with teeth     
powerfully developed1 there is a genus  
named ,ornithostoma from the resemblance
 of its mouth to that of a bird in being
 entirely devoid of teeth4 ,it is       
scarcely possible to distinguish the    
remains of the toothed and toothless    
skeletons except in the dentary         
character of the jaws4 ,there is no     
evidence that the toothless types ever  
possessed a tooth of any sort4 ,they    
were first found in fragments in        

,england in the ,cambridge ,greensand1  
but were afterwards met with in great   
abundance in the ,chalk of ,kansas1     
where the same animals were named       
,pteranodon4 ,a jaw so entirely         
bird-like suggests that the digestive   
organs of ,pterodactyles may in such    
toothless forms at least have been      
characterised by a gizzard1 which is so 
distinctive of ,birds4 ,the absence of  
teeth in the ,great ,ant-eater and some 
other allied ,mammals has transferred   
the function which teeth usually perform
 to the stomach1 one part of which      
becomes greatly thickened and muscular1 
adapting itself to the work which it has
 to perform4 ,it is probable that the   
gizzard may be developed in relation to 
the necessities which food creates1     
since even ,trout1 feeding on the       
shell-fish in some ,irish lochs1 acquire
 such a thickened muscular stomach1 and 
a like modification is recorded in other
 fishes as produced by food4  ,closely  
connected with an animal's habits is the

 protection to the body which is        
afforded by the skin4 ,in ,pterodactyles
 the evidence of the condition of the   
skin is scanty1 and mostly negative4    
,sometimes the dense1 smooth texture of 
the jaw bones indicates a covering like 
the skin of a ,lizard or the hinder part
 of the jaw of a ,bird4 ,some jaws from 
the ,cambridge ,greensand have the bone 
channeled over its surface by minute    
blood vessels which have impressed      
themselves into the bone more easily    
than into its covering4 ,thus in the    
species of ,ornithocheirus distinguished
 as microdon the palate is absolutely   
smooth1 while in the species named      
machaerorhynchus it is marked by        
parallel impressed vascular grooves     
which diverge from the median line4     
,this condition clearly indicates a     
difference in the covering of the bone1 
and that in the latter species the      
covering had fewer blood vessels and    
more horny protection than in the other4
 ,the tissue may not have been of firmer

 consistence than in the palate of      
,mammals4 ,the extremity of the beak is 
often as full of blood vessels as the   
jaw of a ,turtle or ,crocodile4         
,,covering ,,of ,,the ,,body  ,there is 
no trace even in specimens from the     
,solenhofen or ,stonesfield ,slate of   
any covering to the body4 ,there are no 
specimens preserved like mummies1 and   
although the substance of the wings is  
found there is no trace of fur or       
feathers1 bones1 or scales on the skin4 
,the only example in which there is even
 an appearance suggesting feathers is in
 the beautiful ,scaphognathus at ,bonn1 
and upon portions of the wing membrane  
of that specimen are preserved a very   
few small short and apparently tubular  
bodies1 which have a suggestive         
resemblance to the quills of small      
undeveloped feathers4 ,such evidences   
have been diligently sought for4        
,professor ,marsh1 after examining the  
wing membranes of his specimen of       
,rhamphorhynchus from ,solenhofen1      

stated that the wings were partially    
folded and naturally contracted into    
folds1 and that the surface of the      
tissue is marked by delicate striae1    
which might easily be taken at first    
sight for a thin coating of hair4       
,closer investigation proved the        
markings to be minute wrinkles on the   
under surface of the wing membrane4     
,this negative evidence has considerable
 value1 because the ,solenhofen ,slate  
has preserved in the two known examples 
of the bird ,archaeopteryx beautiful    
details of the structure of the larger  
feathers concerned in flight4 ,it has   
preserved many structures far more      
delicate4 ,there is1 therefore1 reason  
for believing that if the skin had      
possessed any covering like one of those
 found in existing vertebrate animals1  
it could scarcely have escaped detection
 in the numerous undisturbed skeletons  
of ,pterodactyles which have been       
examined4  ,the absence of a            
recognisable covering to the skin in a  

fossil state cannot be accepted as      
conclusive evidence of the temperature1 
habits1 or affinities of the animal4    
,although ,mammalia are almost entirely 
clothed with dense hair1 which has never
 been found in a recognisable condition 
in a fossil state in any specimen of    
,tertiary age1 one entire order1 the    
,cetacea1 show in the smooth hairless   
skins of ,whales and ,porpoises that the
 class may part with the typical        
characteristic covering without loss of 
temperature and without intelligible    
cause4 ,that the absence of hair is not 
due to the aquatic conditions of rivers 
or sea is proved by other marine        
,mammals1 like ,seals1 having the skin  
clothed with a dense growth of hair1    
which is not surpassed in any other     
order4 ,the fineness of the growth of   
hair in ,man gives a superficial        
appearance of the skin being imperfectly
 clothed1 and a similar skin in a fossil
 state might give the impression that it
 was devoid of hair4 ,there are many    

,mammals in which the skin is scantily  
clothed with hair as the animal grows   
old4 ,neither the ,elephant nor the     
,armadillo in a fossil state would be   
likely to have the hair preserved1 for  
the growth is thin on the bony shields  
of the living ,armadilloes4 ,yet the    
difficulty need be no more inherent in  
the nature of hair than in that of      
feathers1 since the hair of the ,mammoth
 and ,rhinoceros has been completely    
preserved upon their skins in the       
tundras of ,siberia1 densely clothing   
the body4 ,this may go to show that the 
,pterodactyle possessed a thin covering 
of hair1 or1 more probably1 that hair   
was absent4 ,since ,reptiles are equally
 variable in the clothing of the skin   
with bony or horny plates1 and in       
sometimes having no such protection1 it 
may not appear singular that the skin in
 ,ornithosaurs has hitherto given no    
evidence of a covering4 ,from analogy a 
covering might have been expected2      
feathers of ,birds and hair of ,mammals 

are non-conducting coverings suited to  
arrest the loss of heat4  ,with the     
evidence1 such as it is1 of resemblance 
of ,ornithosaurs to ,birds in some      
features of respiration and flight1 a   
covering to the skin might have been    
expected4 ,yet the covering may not be  
necessary to a high temperature of the  
blood4 ,since ,dr4 ,john ,davy made his 
observations it has been known that the 
temperature of the ,tunny1 above #ij    
deg4 ,fahrenheit1 is as warm as the     
,african scaly ant-eater named the      
,pangolin1 which has the body more amply
 protected by its covering4 ,this       
illustration also shows that hot blood  
may be produced without a four-celled   
heart1 with which it is usually         
associated1 and that even if the skin in
 ,pterodactyles was absolutely naked an 
active life and an abundant supply of   
blood could have given the animal a high
 temperature4  ,the circumstance that in
 several individuals the substance of   
the wing membrane is preserved would    

appear to indicate either that it was   
exceptionally stout when there would    
have been small chance of resisting     
decomposition1 or that its preservation 
is due to a covering which once existed 
of fur or down or other clothing        
substance1 which has proved more durable
 than the skin itself4    ,illustration3
 ,,fig4 #dh4  ,,remains ,,of            
,,dimorphodon ,,from ,,the ,,lias ,,of  
,,lyme   ,,regis    ,,showing ,,the     
,,skull1 ,,neck1 ,,back ,,and ,,some    
,,of ,,the ,,longer ,,bones ,,of ,,the  
,,skeleton    ,from a slab in the       
,british ,museum 7,natural ,history7    
,,chapter ,,xiii  ,,ancient             
,,ornithosaurs ,,from ,,the ,,lias      
,cuvier's discourse on the revolutions  
of the ,earth made the ,pterodactyle    
known to ,english readers early in the  
nineteenth century4 ,dr4 ,buckland1 the 
distinguished professor of ,geology at  
,oxford1 discovered in #ahbi a far      
larger specimen in the ,lias of ,lyme   
,regis1 and it became known by a figure 

published by the ,geological ,society1  
and by the description in his famous    
,bridgewater ,treatise1 p4 #afd4 ,this  
animal was tantalising in imperfect     
preservation4 ,the bones were scattered 
in the clay1 so as to give no idea of   
the animal's aspect4 ,knowledge of its  
limbs and body has been gradually       
acquired2 and now1 for some years1 the  
tail and most parts of the skeleton have
 been well known in this oldest and most
 interesting ,british ,pterodactyle4    
,sir ,richard ,owen after some time     
separated the fossil as a distinct      
genus1 named ,dimorphodon2 for it was in
 many ways unlike the ,pterodactyles    
described from ,bavaria4 ,the name      
,dimorphodon indicated the two distinct 
kinds of teeth in the jaws1 a character 
which is still unparalleled among       
,pterodactyles of newer age4 ,there are 
a few large pointed1 piercing and       
tearing teeth in the front of the jaws1 
with smaller teeth further back1 placed 
among the tearing teeth in the upper    

jaw2 while in the lower jaw the small   
teeth are continuous1 close-set1 and    
form a fine cutting edge like a saw4    
,illustration3 ,,fig4 #di4  ,,left      
,,side ,,of ,,dimorphodon 7,,restored7  
,,at ,,rest  ,the ,dimorphodon has a    
short beak1 a deep head1 and deep lower 
jaw1 which is overlapped by the cheek   
bones4 ,the side of the head is occupied
 by four vacuities1 separated by narrow 
bars of bone4 ,first1 in front1 is the  
immense opening for the nostril1        
triangular in form1 with the long upper 
side following the rounded curve of the 
face4 ,a large triangular opening       
intervenes between the nose hole and the
 eye hole1 scarcely smaller than the    
former1 but much larger than the orbit  
of the eye4 ,the eye hole is shaped like
 a kite or inverted pear4 ,further back 
still is a narrower vertical opening    
known as the lateral or inferior        
temporal vacuity4 ,the back of the head 
is badly preserved4 ,the two principal  
skulls differ in depth1 probably from   

the strains under which they were       
pressed flat in the clay4 ,a singular   
detail of structure is found in the     
extremity of the lower jaw1 which is    
turned slightly downward1 and terminates
 in a short toothless point4 ,the head  
of ,dimorphodon is about eight inches   
long4    ,illustration3 ,,fig4 #ej4     
,,dimorphodon ,,macronyx   ,,restored   
,,form ,,of ,,the ,,animal  ,the neck   
bones are of suitable stoutness and     
width to support the head4 ,the bones   
are yoked together by strong processes4 
,the neck was about #f inches long1 did 
not include more than seven bones1 and  
appeared short owing only to the depth  
and size of the head4 ,the length of the
 backbone which supported the ribs was  
also about #f inches4 ,its joints are   
remarkably short when compared with     
those of the neck4 ,the tail is about   
#bj inches long4  ,the extreme length of
 the animal from the tip of the nose to 
the end of the tail may have been #c    
feet #d inches1 supposing it to have    

walked on all fours in the manner of a  
,reptile or ,mammal4 ,this may have been
 a common position1 but ,dimorphodon may
 probably also have been a biped4       
,before #ahge1 when the first           
restoration appeared in the ,illustrated
 ,london ,news1 the legs had been       
regarded as too short to have supported 
the animal1 standing upon its hind      
limbs4 ,they are here seen to be well   
adapted for such a purpose4 ,the upper  
leg bone is #c-a_/d inches long1 the    
lower leg bone is #d-a_/b inches long1  
and the singularly strong instep bones  
are firmly packed together side by side 
as in a leaping or jumping ,mammal1 and 
measure #a-a_/b inches in length4       
,dimorphodon differs from several other 
,pterodactyles in having the hind limb  
provided with a fifth outermost short   
instep bone1 to which two toe bones are 
attached4 ,these bones are elongated in 
a way that may be compared1 on a small  
scale1 with the elongation of the wing  
finger in the fore limb4 ,the digit was 

manifestly used in the same way as the  
wing finger1 in partial support of a    
flying membrane1 though its direction   
may have been upward and outward1 rather
 than inward4 ,there is no evidence of a
 pulley joint between the metatarsal and
 the adjacent phalange4  ,the height of 
the ,dimorphodon1 standing on its hind  
legs in the position of a ,bird1 with   
the wings folded upon the body in the   
manner of a ,bird1 was about #bj inches4
 ,an ungainly1 ill-balanced animal in   
aspect1 but not more so than many       
big-headed birds1 and probably capable  
of resting upon the instep bones as many
 birds do4 ,the chief point of variation
 from the ,pterodactyle wing is in the  
relative length of the metacarpus in    
,dimorphodon4 ,it is shorter than the   
other bones in the wing1 never exceeding
 #a-a_/b inches4 ,the total length of   
all the arm bones down to the point     
where the metacarpus might have touched 
the ground1 or where the wing finger is 
bent upon it1 is about #i inches1 which 

gives a length of less than #f inches   
below the upper arm bone4 ,the four     
bones of the wing finger measure1 from  
the point where the first bone bends    
upon the metacarpus1 less than #ah      
inches4 ,so that the wings could only   
have been carried in the manner of the  
wings of a ,bat1 folded at the side and 
directed obliquely over the back when   
the animal moved on all fours4 ,its body
 would appear to have been raised high  
above the ground1 in a manner almost    
unparalleled in ,reptiles1 and          
comparable to ,birds and ,mammals4      
,dimorphodon is to be imagined in full  
flight1 with the body extended like that
 of a ,bird1 when the wings would have  
had a spread from side to side of about 
#d feet #d inches4 ,as in other animals 
of this group1 the three claws on the   
front feet are larger than the similar  
four claws on the hind feet2 as though  
the fingers might have functions in     
grasping prey1 which were not shared by 
the toes4    ,illustration3 ,,fig4 #ea4 

,,dimorphodon ,,macronyx ,,walking ,,as 
,a ,,quadruped   ,,restoration ,,of     
,,the ,,skeleton  ,the restorations give
 faithful pictures of the skeleton1 and 
the form of the body is built upon the  
indications of muscular structure seen  
in the bones4  ,a second ,english       
,pterodactyle is found in the ,upper    
,lias of ,whitby4 ,it is only known from
 an imperfect skull1 published in #ahhh4
 ,it has the great advantage of         
preserving the bones in their natural   
relations to each other1 and with a     
length of head probably similar to      
,dimorphodon shows that the depth at the
 back of the eye was much less2 and the 
skull wants the arched contour of face  
seen in ,dimorphodon4 ,the head has the 
same four lateral vacuities1 but the    
nostril is relatively small and         
elongated1 extending partly above the   
oval antorbital opening1 which was      
larger4 ,there is thus a difference of  
proportion1 but it is precisely such as 
might result from the species having the

 skull flatter4 ,the head is easily     
distinguished by the small nostril1     
which is smaller than the orbit of the  
eye4 ,the animal is referred to another 
genus4 ,the quadrate bones which give   
attachment to the lower jaw send a      
process inward to meet the bones of the 
palate1 which differ somewhat from the  
usual condition4 ,two bony rods extend  
from the quadrate bones backward and    
upward to the sphenoid1 and two more    
slender bones extend from the quadrate  
bones forward1 and converge in a        
,v-shape1 to define the division between
 the openings of the nostrils on the    
palate4 ,the ,v-shaped bone in front is 
called the vomer1 while the hinder part 
is called pterygoid4 ,the bones that    
extend backward to the sphenoid are not 
easily identified4 ,this animal is one  
of the most interesting of              
,pterodactyles from the very reptilian  
character exhibited in the back of the  
head1 which appears to be different from
 other specimens1 which are more like a 

bird in that region4 ,yet underneath    
this reptilian aspect1 with the bony bar
 at the side of the temporal region of  
the head formed by the squamosal and    
quadrate bones1 defining the two        
temporal vacuities as in ,reptiles1 a   
mould is preserved of the cavity once   
occupied by the brain1 showing the chief
 details of structure of that organ1 and
 proving that in so far as it departs   
from the brain of a ,bird it appears to 
resemble the brain of a ,mammal1 and is 
unlike the brain of a ,reptile4  ,the   
,pterodactyles from the ,lias of        
,germany are similar to the ,english    
types1 in so far as they can be         
compared4 ,in #ahgh ,i had the          
opportunity of studying those which were
 preserved in the ,castle at ,banz1     
which ,professor ,andreas ,wagner1 in   
#ahfj1 referred to the new genus        
,dorygnathus4 ,the skull is unknown1 but
 the lower jaw1 #f-a_/b inches long1 is 
less than #b-a_/b inches wide at the    
articulation with the quadrate bone in  

the skull4 ,the depth of the lower jaw  
does not exceed #a_/d inch1 so that it  
is in marked contrast to ,buckland's    
,dimorphodon4 ,the symphysis1 which     
completely blends the rami of the jaw1  
is short4 ,as far as it extends it      
contains large tearing teeth1 followed  
by smaller teeth behind1 like those of  
,dimorphodon4 ,but this ,german fossil  
appears to differ from the ,english type
 in having the front of the lower jaw1  
for about #c_/d inch1 compressed from   
side to side into a sharp blade or      
spear1 more marked than in any other    
,pterodactyle1 and directed upward      
instead of downward as in ,dimorphodon4 
,nearly all the measurements in the     
skeleton are practically identical with 
those of the ,english ,dimorphodon1 and 
extend to the jaw1 humerus1 ulna and    
radius1 wing metacarpal1 first phalange 
of the wing finger4 ,the principal bones
 of the hind limb appear to be a little 
shorter2 but the scapula and coracoid   
are slightly larger4 ,all these bones   

are so similar in form to ,dimorphodon  
that they could not be separated from   
the ,lyme ,regis species1 if they were  
found in the same locality4             
,illustration3 ,,fig4 #eb4              
,,dimorphodon ,,macronyx ,,walking ,,as 
,a ,,biped   ,based chiefly on remains  
in the ,british ,museum                 
,illustration3 ,,fig4 #ec4  ,,lower     
,,jaw ,,of ,,dorygnathus ,,seen ,,from  
,,below    ,from the ,lower ,lias of    
,germany1 showing the spear in front of 
the   tooth sockets  ,just as the ,upper
 ,lias in ,england has yielded a second 
,pterodactyle1 so the ,upper ,lias in   
,germany has yielded a skeleton1 to     
which ,felix ,plieninger1 in #ahid1 gave
 the name ,campylognathus4 ,it is an    
instructive skeleton1 with the head much
 smaller than in ,dimorphodon1 being    
less than #f inches long1 but1          
unfortunately1 broken and disturbed4 ,a 
lower jaw gives the length #d-a_/b      
inches4 ,like the other ,pterodactyles  
from the ,lias1 it has the extremity of 

the beak toothless1 with larger teeth in
 the region of the symphysis in front   
and smaller teeth behind4 ,the jaw is   
deeper than in the ,banz specimen from  
the ,lower ,lias1 but not so deep as in 
,dimorphodon4 ,the teeth of the upper   
jaw vary in size1 and there appears to  
be an exceptionally large tooth in the  
position of the ,mammalian canine at the
 junction of the bones named maxillary  
and intermaxillary4  ,the nasal opening 
is small and elongated1 as in the       
,english specimen from ,whitby4 ,as in  
that type there is little or no         
indication of the convex contour of the 
face seen in ,dimorphodon4  ,the neck   
does not appear to be preserved4 ,in the
 back the vertebrae are about #c_/aj    
inch long1 so that twelve1 which is the 
usual number1 would only occupy a length
 of a little more than #c-a_/b inches4  
,the tail is elongated like that of     
,dimorphodon1 and bordered in the same  
way by ossified ligaments4 ,there are   
thirty-five tail vertebrae4 ,those which

 immediately follow the pelvis are      
short1 like the vertebrae of the back4  
,but they soon elongate1 and reach a    
maximum length of nearly #a-a_/b inches 
at the eighth1 and then gradually       
diminish till the last scarcely exceeds 
#a_/h inch in length4 ,the length of the
 tail is about #bb inches2 this appears 
to be an inch or two longer than in     
,dimorphodon4 ,the longest rib measures 
#b-a_/b inches1 and the shortest #b     
inches4 ,these ribs probably were       
connected with the sternum1 which is    
imperfectly preserved4    ,illustration3
 ,,fig4 #ed4  ,,dimorphodon ,,macronyx  
,,showing ,,the ,,maximum ,,spread ,,of 
,,the ,,wing ,,membranes  ,the bones of 
the limbs have about the same length as 
those of ,dimorphodon1 so far as they   
can be compared1 except that the ulna   
and radius are shorter4 ,the wing       
metacarpal is of about the same length1 
but the first phalange of the wing      
finger measures #f-a_/d inches1 the     
second is about #h-a_/d inches1 the     

third #f-a_/b inches1 and the fourth    
#d-c_/d inches2 so that the total length
 of the wing finger was about half an   
inch short of #b feet4 ,one character   
especially deserves attention in the    
apparent successive elongation of the   
first three phalanges in the wing finger
 in ,dimorphodon4 ,the third phalange is
 the longest in the only specimen in    
which the finger bones are all          
preserved4 ,usually the first phalange  
is much longer than the second1 so that 
it is a further point of interest to    
find that this ,german type shares with 
,dimorphodon a character of the wing    
finger which distinguishes both from    
some members of the group by its short  
first phalange4    ,illustration3 ,,fig4
 #ee4  ,,the ,,left ,,side ,,of ,,the   
,,pelvis ,,of ,,dimorphodon   ,,showing 
,,the ,,two ,,prepubic ,,bones  ,the    
pelvis is exceptionally strong in       
,campylognathus1 and although it is     
crushed the bones manifestly met at the 
base of the ischium1 while the pubic    

bones were separated from each other in 
front4 ,the bones of the hind limb are  
altogether shorter in the ,german fossil
 than in ,dimorphodon1 especially in the
 tibia2 but the structure of the        
metatarsus is just the same1 even to the
 short fifth metatarsal with its two    
digits1 only those bones are extremely  
short1 instead of being elongated as in 
,dimorphodon4 ,it is therefore          
convenient1 from the different          
proportions of the body1 that           
,campylognathus may be separated from   
,dimorphodon2 but so much as is         
preserved of the ,english specimen from 
the ,upper ,lias of ,whitby rather      
favours the belief that our species     
should also be referred to              
,campylognathus1 which had not been     
figured when the ,whitby skull was      
referred to ,scaphognathus by ,mr4      
,newton4 ,it may be doubtful whether    
there is sufficient evidence to         
establish the distinctness of the other 
,german genus ,dorygnathus1 though it   

may be retained pending further         
knowledge4  ,in these characters are    
grounds for placing the ,lias           
,pterodactyles in a distinct family1 the
 ,dimorphodontidae1 as was suggested in 
#ahgj4 ,this evidence is found in the   
five metatarsal bones1 of which four are
 in close contact1 the middle two being 
slightly the longest1 so as to present  
the general aspect of the corresponding 
bones in a ,mammal rather than a ,bird4 
,secondly1 the very slender fibula1     
prolonged down the length of the shin   
bone1 which ends in a rounded pulley    
like the corresponding bone of a ,bird4 
,thirdly1 the great elongation of the   
third wing phalange4 ,fourthly1 the     
prolongation of the coracoid bone beyond
 the articulation for the humerus1 as in
 a ,bird4 ,and the toothless1           
spear-shaped beak1 and jaw with large   
teeth in front and small teeth behind1  
are also distinctive characters4        
,,chapter ,,xiv  ,,ornithosaurs ,,from  
,,the ,,middle ,,secondary ,,rocks      

,,rhamphocephalus  ,,the ,stonesfield   
,slate in ,england1 which corresponds in
 age with the lower part of the ,great  
or ,bath ,oolite1 yields many evidences 
of terrestrial life--land plants1       
insects1 and mammals--preserved in a    
marine deposit4 ,a number of isolated   
bones have been found of ,pterodactyles1
 some of them indicating animals of     
considerable size and strength4 ,the    
nature of the limestone was unfavourable
 to the preservation of soft wing       
membranes1 or even to the bones         
remaining in natural association4 ,very 
little is known of the head of          
,rhamphocephalus4 ,one imperfect        
specimen shows a long temporal region   
which is wide1 and a very narrow        
interspace between the orbits2 with a   
long face1 indicated by the extension of
 narrow nasal bones4 ,the lower jaw has 
an edentulous beak or spear in front1   
which is compressed from side to side in
 the manner of the ,liassic forms1 but  
turned upward slightly1 as in           

,dorygnathus or ,campylognathus4 ,behind
 this extremity are sharp1 tall teeth1  
few in number1 which somewhat diminish  
in size as they extend backward1 and do 
not suddenly change to smaller series1  
as in the ,lias genera4 ,a few small    
vertebrae have been found1 indicating   
the neck and back4 ,the sacrum consists 
of five vertebrae4 ,one small example   
has a length of only an inch4 ,it is a  
little narrower behind than in front1   
and would be consistent with the animal 
having had a long tail1 which ,i believe
 to have been present1 although ,i have 
not seen any caudal vertebrae4 ,the     
early ribs are like the early ribs of a 
,crocodile or ,bird in the well-marked  
double articulation4 ,the later ribs    
appear to have but one head4 ,v-shaped  
abdominal ribs are preserved4 ,much of  
the animal is unknown4 ,the coracoid    
seems to have been directed forward1    
and1 as in a bird1 it is #b-a_/b inches 
long4 ,the humerus is #c-a_/b inches    
long1 and the fore-arm measured #f      

inches1 so that it was relatively longer
 than in ,dimorphodon4 ,the metacarpus  
is #a-c_/d inches long4 ,the wing finger
 was exceptionally long and strong4     
,professor ,huxley gave its length at   
#bi inches4 ,my own studies lead to the 
conclusion that the first finger bone of
 the wing was the shorter1 and that     
although they did not differ greatly in 
length1 the second was probably the     
longest1 as in ,campylognathus4         
,professor ,huxley makes the second and 
third phalanges #g-c_/d inches long1 and
 the first only about #c_/h inch        
shorter1 while the fourth phalange is   
#f-a_/b inches4 ,these measurements are 
based upon some specimens in the ,oxford
 ,university ,museum4 ,there is only one
 first phalange which has a length of   
#g-c_/d inches4 ,the others are between 
#e and #f inches1 or but little exceed  
#d inches2 so that as all the fourth    
phalanges which are known have a length 
of #f-a_/b inches1 it is possible that  
the normal length of the first phalange 

in the larger species was #e-a_/b       
inches4 ,the largest of the phalanges   
which may be classed as second or third 
is #h-a_/b inches1 and that1 ,i suppose1
 may have been associated with the      
#g-c_/d inches first phalange4 ,but the 
other bones which could have had this   
position all measure #e-a_/b and #g-c_/d
 inches4 ,the three species indicated by
 finger bones may have had the          
measurements3--       ,phalanges of the 
wing finger    _/                       
,i4        ,,ii4      ,,iii4      ,,iv4 
#g-c_/d     #h-a_/b      #g8            
#f-a_/b          length of each bone    
#e-a_/b     #g-c_/d     #e-a_/b         
#d-a_/b8             in inches4         
#d-a_/b     ----      ----       ----   
,the femur is represented by many       
examples--one #c-c_/d inches long1 and  
others less than #c inches long         
7#b-i_/aj74 ,in ,campylognathus1 which  
has so much in common with the jaw and  
the wing bones in size1 the upper leg   
bone is #b-h_/aj inches4 ,therefore if  

we assign the larger femur to the larger
 wing1 the femur will be relatively     
longer in all species of                
,rhamphocephalus than in                
,campylognathus4 ,only one example of a 
tibia is preserved4 ,it is #c-a_/b      
inches long1 or only #a_/aj inch shorter
 than the bone in ,campylognathus1 which
 has the femur #b-h_/aj inches1 so that 
,i refer the tibia of ,rhamphocephalus  
to the species which has the            
intermediate length of wing4 ,these     
coincidences with ,campylognathus       
establish a close affinity1 and may     
raise the question whether the ,upper   
,lias species may not be included in the
 ,stonesfield ,slate genus              
,rhamphocephalus4  ,the late ,professor 
,phillips1 in his ,geology of ,oxford1  
attempted a restoration of the          
,stonesfield ,ornithosaur1 and produced 
a picturesque effect 7p4 #afd72 but no  
restoration is possible without such    
attention to the proportions of the     
bones as we have indicated4   ,,oxford  

,,clay  ,a few bones of flying reptiles 
have been found in the ,lower ,oxford   
,clay near ,peterborough1 and others in 
the ,upper ,oxford ,clay at ,st4 ,ives1 
in ,huntingdonshire4 ,a single tail     
vertebra from the ,middle ,oxford ,clay1
 near ,oxford1 long since came under my 
own notice1 and shows that these animals
 belong to a long-tailed type like      
,campylognathus4 ,the cervical vertebrae
 are remarkable for being scarcely      
longer than the dorsal vertebrae2 and   
the dorsal are at least half as long    
again as is usual1 having rather the    
proportion of bones in the back of a    
crocodile4   ,,lithographic ,,slate     
,long-tailed ,pterodactyles are         
beautifully preserved in the            
,lithographic ,limestone of the south of
 ,bavaria1 at ,solenhofen1 and the      
quarries in its neighbourhood1 often    
with the skeleton or a large part of it 
flattened out in the plane of bedding of
 the rock4 ,fine skeletons are preserved
 in the superb museum at ,munich1 at    

,heidelberg1 ,bonn1 ,haarlem1 and       
,london1 and are all referred to the    
genus ,rhamphorhynchus or to            
,scaphognathus4 ,it is a type with      
powerfully developed wings and a long1  
stiff tail1 very similar to that of     
,dimorphodon1 so that some naturalists  
refer both to the same family4 ,there is
 some resemblance4  ,the type which is  
most like ,dimorphodon is the celebrated
 fossil at ,bonn1 sometimes called      
,pterodactylus crassirostris1 which in a
 restored form1 with a short tail1 has  
been reproduced in many text-books4 ,no 
tail is preserved in the slab1 and ,i   
ventured to give the animal a tail for  
the first time in a restoration 7p4     
#afc7 published by the ,illustrated     
,london ,news in #ahge1 which           
accompanied a report of a ,royal        
,institution lecture4 ,afterwards1 in   
#ahhb1 ,professor ,zittel1 of ,munich1  
published the same conclusion4 ,the     
reason for restoring the tail was that  
the animal had the head constructed in  

the same way as ,pterodactyles with a   
long tail1 and showed differences from  
types in which the tail is short2 and   
there is no known short-tailed          
,pterodactyle1 with wrist and hand      
bones1 such as characterise this animal4
 ,the side of the face has a general    
resemblance to the ,pterodactyles from  
the ,lias1 for although the framework is
 firmer1 the four apertures in the head 
are similarly placed4 ,the nostril is   
rather small and elongated1 and ascends 
over the larger antorbital vacuity4 ,the
 orbit for the eye is the largest       
opening in the head1 so that these three
 apertures successively increase in     
size1 and are followed by the vertically
 elongated post-orbital vacuity4 ,the   
teeth are widely spaced apart1 and those
 in the skull extend some distance      
backward to the end of the maxillary    
bone4 ,there are few teeth in the lower 
jaw1 and they correspond to the large   
anterior teeth of ,dimorphodon1 there   
being no teeth behind the nasal opening4

 ,the lower jaw is straight1 and the    
extremities of the jaws met when the    
mouth was closed4 ,the breast bone does 
not show the keel which is so remarkable
 in ,rhamphorhynchus1 which may be      
attributed to its under side being      
exposed1 so as to exhibit the pneumatic 
foramina4  ,the ribs have double heads1 
more like those of a ,crocodile in the  
region of the back than is the case with
 the bird-like ribs from ,stonesfield4  
,the second joint in the wing finger may
 be longer than the first--a character  
which would tend to the association of  
this ,pterodactyle with species from the
 ,lias2 a relation to which attention   
was first drawn by ,mr4 ,e4 ,t4 ,newton1
 who described the ,whitby skull4  ,the 
,pterodactyles from the ,solenhofen     
,slate which possess long tails have a  
series of characters which show affinity
 with the other long-tailed types4 ,the 
jaws are much more slender4 ,the orbit  
of the eye in ,rhamphorhynchus is       
enormously large1 and placed vertically 

above the articulation for the lower    
jaw4 ,immediately in front of the eye   
are two small and elongated openings1   
the hinder of which1 known as the       
antorbital vacuity1 is often slightly   
smaller than the nostril1 which is      
placed in the middle length of the head1
 or a little further back1 giving a long
 dagger-shaped jaw1 which terminates in 
a toothless spear4 ,the lower jaw has a 
corresponding sharp extremity4 ,the     
teeth are directed forward in a way that
 is quite exceptional4 ,notwithstanding 
the massiveness and elongation of the   
neck vertebrae1 which are nearly twice  
as long as those of the back1 the neck  
is sometimes only about half the length 
of the skull4  ,all these long-tailed   
species from the ,lithographic ,stone   
agree in having the sternum broad1 with 
a long strong keel1 extending far       
forward4 ,the coracoid bones extend     
outward like those of a ,crocodile1 so  
as to widen the chest cavity instead of 
being carried forward as the bones are  

in ,birds4 ,these bones in this animal  
were attached to the anterior extremity 
of the sternum1 so that the keel        
extended in advance of the articulation 
as in other ,pterodactyles4 ,the breadth
 of the sternum shows that1 as in       
,mammals1 the fore part of the body must
 have been fully twice the width of the 
region of the hip-girdle1 where the     
slenderer hind limbs were attached4 ,the
 length of the fore limb was enormous1  
for although the head suggests an       
immense length relatively to the body1  
nearly equal to neck and back together1 
the head is not more than a third of the
 length of the wing bones4 ,the wing    
bones are remarkable for the short      
powerful humerus with an expanded radial
 crest1 which is fully equal in width to
 half the length of the bone4 ,another  
character is the extreme shortness of   
the metacarpus1 usually associated with 
immense strength of the wing metacarpal 
bone4  ,the hind limbs are relatively   
small and relatively short4 ,the femur  

is usually shorter than the humerus1 and
 the tibia is much shorter than the     
ulna4 ,the bones of the instep1 instead 
of being held together firmly as in the 
,lias genera1 diverge from each other1  
widening out1 though it often happens   
that four of the five metatarsals differ
 but little in length4 ,the fifth digit 
is always shorter4  ,the hip-girdle of  
bones differs chiefly from other types  
in the way in which those bones1 which  
have sometimes been likened to the      
marsupial bones1 are conditioned4 ,they 
may be a pair of triangular bones which 
meet in the middle line1 so that there  
is an outer angle like the arm of a     
capital ,y4 ,sometimes these triangular 
bones are blended into a curved1        
bow-shaped arch1 which in several       
specimens appears to extend forward from
 near the place of articulation of the  
femur4 ,this is seen in fossil skeletons
 at ,heidelberg and ,munich4 ,it is     
possible that this position is an       
accident of preservation1 and that the  

prepubic bones are really attached to   
the lower border of the pubic bones4    
,immense as the length of the tail      
appears to be1 exceeding the skull and  
remainder of the vertebral column1 it   
falls far short of the combined length  
of the phalanges of the wing finger4    
,the power of flight was manifestly     
greater in ,rhamphorhynchus than in     
other members of the group1 and all the 
modifications of the skeleton tend      
towards adaptation of the animals for   
flying4 ,the most remarkable            
modification of structure at the        
extremity of the tail was made known by 
,professor ,marsh in a vertical1        
leaf-like expansion in this genus1 which
 had not previously been observed 7p4   
#afa74 ,the vertebrae go on steadily    
diminishing in length in the usual way1 
and then the ossified structures which  
bordered the tail bones and run parallel
 with the vertebrae in all the          
,rhamphorhynchus family1 suddenly       
diverge downward and upward at right    

angles to the vertebrae1 forming a      
vertical crest above and a corresponding
 keel below2 and between these          
structures1 which are identified with   
the neural spines and chevron bones of  
ordinary vertebrae1 the membrane        
extends1 giving the extremity of the    
tail a rudder-like feature1 which1 from 
knowledge of the construction of the    
tail of a child's kite1 may well be     
thought to have had influence in        
directing and steadying the animal's    
movements4 ,there are many minor        
features in the shoulder-girdle1 which  
show that the coracoid1 for example1 was
 becoming unlike that bone in the ,lias1
 though it still continues to have a    
bony union with the elongated           
shoulder-blade of the back4             
,illustration3 ,,fig4 #ef4              
,,restoration ,,of ,,the ,,skeleton ,,of
   ,,rhamphorhynchus ,,phyllurus        
,from the ,solenhofen ,slate1 partly    
based upon the skeleton   with the wing 
membranes preserved    ,illustration3   

,,fig4 #eg4  ,,restoration ,,of ,,the   
,,skeleton ,,of ,,scaphognathus         
,,crassirostris    ,published in the    
,illustrated ,london ,news in #ahge4 ,in
 which a tail is   shown on the evidence
 of the structure of the head and hand  
,illustration3 ,,fig4 #eh4  ,,six       
,,restorations      #a4 ,ramphocephalus4
 ,stonesfield ,slate4 ,john ,phillips1  
#ahga     #b4 ,rhamphorhynchus4 ,o4 ,c4 
,marsh1 #ahhb     #c4 ,rhamphorhynchus4 
,v4 ,zittel1 #ahhb     #d4              
,ornithostoma4 ,williston1 #ahig     #e4
 ,dimorphodon4 ,buckland1 #ahcf4 ,tail  
then unknown     #f4 ,ornithocheirus4   
,h4 ,g4 ,seeley1 #ahfe  ,the great      
,german delineator of these animals1    
,von ,meyer1 admitted six different     
species4 ,mr4 ,newton and ,mr4 ,lydekker
 diminish the number to four4 ,it is not
 easy to determine these differences1 or
 to say how far the differences observed
 in the bones characterise species or   
genera4 ,it is certain that there is one
 remarkable difference from other and   

older ,pterodactyles1 in that the last  
or fourth bone in the wing finger is    
usually slightly longer than the third  
bone1 which precedes it4 ,there is a    
certain variability in the specimens    
which makes discussion of their         
characters difficult1 and has led to    
some forms being regarded as varieties1 
while others1 of which less material is 
available1 are classed as species4 ,i am
 disposed to say that some of the       
confusion may have resulted from        
specimens being wrongly named4 ,thus1   
there is a ,rhamphorhynchus called      
curtimanus1 or the form with the short  
hand4 ,it is represented by two types4  
,one of these appears to have the       
humerus short1 the ulna and radius long1
 and the finger bones long2 the other   
has the humerus longer1 the ulna much   
shorter1 and the finger bones shorter4  
,they are clearly different species1 but
 the second variety agrees in almost    
every detail with a species named       
hirundinaceus1 the swallow-like         

,rhamphorhynchus4 ,this identification  
shows1 not that the latter is a bad     
species1 but that curtimanus is a       
distinct species which had sometimes    
been confounded with the other4 ,while  
most of these specimens show a small but
 steady decrease in the length of the   
several wing finger bones1 the species  
called ,gemmingi has the first three    
bones absolutely equal and shorter than 
in the species curtimanus1 longimanus1  
or hirundinaceus4 ,in the same way1 on  
the evidence of facts1 ,i find myself   
unable to join in discarding ,professor 
,marsh's species phyllurus1 on account  
of the different proportions of its limb
 bones4 ,the humerus1 metacarpus1 and   
third phalange of the wing finger in    
,rhamphorhynchus phyllurus are          
exceptionally short as compared with    
other species4 ,everyone agrees that the
 species called longicaudus is a        
distinct one1 so that it is chiefly in  
slight differences in the proportions of
 constituent parts of the skeleton that 

the types of the ,rhamphorhynchus are   
distinguished from each other4 ,i cannot
 quite concur with either ,professor    
,zittel 7,fig4 #eh1 #c7 or ,professor   
,marsh 7,fig4 #eh1 #b7 in the expansion 
which they give to the wing membrane in 
their restorations2 for although        
,professor ,zittel represents the tail  
as free from the hind legs1 while       
,professor ,marsh connects them         
together1 they both concur in carrying  
the wing membrane from the tip of the   
wing finger down to the extremity of the
 ankle joint4 ,i should have preferred  
to carry it no further down the body    
than the lower part of the back1 there  
being no fossil evidence in favour of   
this extension so far as specimens have 
been described4 ,neither the membranous 
wings figured by ,zittel nor by ,marsh  
would warrant so much body membrane as  
the ,rhamphorhynchus has been credited  
with4 ,i have based my restoration 7p4  
#afa7 of the skeleton chiefly on        
,rhamphorhynchus phyllurus4   ,,the     

,,short-,,tailed ,,types  ,the          
,pterodactylia are less variable2 and   
the variation among the species is      
chiefly confined to relative length of  
the head1 length of the neck1 and the   
height of the body above the ground4    
,the tail is always so short as to be   
inappreciable4 ,many of the specimens   
are fragmentary1 and the characters of  
the group are not easily determined     
without careful comparisons and         
measurements4 ,the bones of the fore    
limb and wing finger are less stout than
 in the ,rhamphorhynchus type1 while the
 femur is generally a little longer than
 the humerus1 and the wing finger is    
short in comparison with its condition  
in ,rhamphorhynchus4 ,these short-tailed
 ,pterodactyles give the impression of  
being active little animals1 having very
 much the aspect of birds1 upon four    
legs or two4 ,the neck is about as long 
as the lower jaw1 the antorbital vacuity
 in the head is imperfectly separated   
from the much larger nasal opening1 the 

orbit of the eye is large and far back1 
the teeth are entirely in front of the  
nasal aperture1 and the post-orbital    
vacuity is minute and inconspicuous4    
,the sternum is much wider than long1   
and no specimens give evidence of a     
manubrium4 ,the finger bones            
progressively decrease in length4 ,the  
prepubic bones have a partially expanded
 fan-like form1 and never show the      
triradiate shape1 and are never         
anchylosed4 ,about fifteen different    
kinds of ,pterodactyles have been       
described from the ,solenhofen ,slate1  
mostly referred to the genus            
,pterodactylus1 which comprises forms   
with a large head and long snout4 ,some 
have been placed in a genus             
7,ornithocephalus1 or ,ptenodracon7 in  
which the head is relatively short4 ,the
 majority of the species are relatively 
small4 ,the skull in ,ornithocephalus   
brevirostris is only #a inch long1 and  
the animal could not have stood more    
than #a-a_/b inches to its back standing

 on all fours1 and but little over      
#b-a_/b inches standing as a biped1 on  
the hind limbs4  ,a restoration of the  
species called ,pterodactylus           
scolopaciceps1 published in #ahge in the
 ,illustrated ,london ,news in the      
position of a quadruped1 shows an animal
 a little larger1 with a body #b-a_/b   
inches high and #f to #g inches long1   
with the wing finger #d-a_/b inches     
long4 ,larger animals occur in the same 
deposit1 and in one named ,pterodactylus
 grandis the leg bones are a foot long2 
and such an animal may have been nearly 
a foot in height to its back1 standing  
as a quadruped1 though most of these    
animals had the neck flexible and       
capable of being raised like the neck of
 a ,goose or a ,deer 7p4 #cj71 and bent 
down like a ,duck's when feeding4       
,illustration3 ,,fig4 #ei4              
,,restoration ,,of ,,the ,,skeleton ,,of
 ,,ptenodracon   ,,brevirostris    ,from
 the ,solenhofen ,slate4 ,the fourth    
joint of the wing finger appears   to be

 lost and has not been restored in the  
figure4 7,natural size7  ,the type of   
the genus ,pterodactylus is the form    
originally described by ,cuvier as      
,pterodactylus longirostris 7p4 #bh74   
,it is also known as ,p4 antiquus1 that 
name having been given by a ,german     
naturalist after ,cuvier had invented   
the genus1 and before he had named the  
species4 ,there are some remarkable     
features in which ,cuvier's animal is   
distinct from others which have been    
referred to the same genus4 ,thus the   
head is #d-a_/b inches long1 while the  
entire length of the backbone to the    
extremity of the tail is only #f-a_/b   
inches1 and one vertebra in the neck is 
at least as long as six in the back1 so 
that the animal has the greater part of 
its length in the head and neck1        
although the neck includes so few       
vertebrae4 ,nearly all the teeth--which 
are few in number1 short and broad1 not 
exceeding a dozen in either jaw--are    
limited to the front part of the beak1  

and do not extend anywhere near the     
nasal vacuity4 ,this is not the case    
with all4  ,in the species named ,p4    
,kochi1 which ,i have regarded as the   
type of a distinct genus1 there are     
large teeth in the front of the jaw     
corresponding to those of               
,pterodactylus1 and behind these a      
smaller series of teeth extending back  
under the nostril1 which approaches     
close to the orbit of the eye1 without  
any indication of a separate antorbital 
vacuity4 ,on those characters the genus 
,diopecephalus was defined4 ,it is      
closely allied to ,pterodactylus2 both  
agree in having the ilium prolonged     
forward more than twice as far as it is 
carried backward1 the anterior process  
covering about half a dozen vertebrae1  
as in ,pterodactylus longirostris4 ,a   
great many different types have been    
referred to ,pterodactylus ,kochi1 and  
it is probable that they may eventually 
be distinguished from each other4 ,the  
species in which the upper borders of   

the orbits approximate could be         
separated from those in which the       
frontal interspace is wider4            
,illustration3 ,,fig4 #fj4              
,,cycnorhamphus ,,suevicus ,,from ,,the 
,,solenhofen   ,,slate ,,showing ,,the  
,,scattered ,,position ,,of ,,the       
,,bones    ,original in the ,museum at  
,tuebingen    ,illustration3 ,,fig4 #fa4
  ,,cycnorhamphus ,,suevicus            
,,restoration ,,showing ,,the ,,form    
,,of ,,the ,,body ,,and ,,the ,,wing    
,,membranes  ,it is a remarkable feature
 in these animals that the middle bones 
of the foot1 termed instep bones or     
metatarsals1 are usually close together1
 so that the toes diverge from a narrow 
breadth1 as in ,p4 longirostris1 ,p4    
,kochi1 and other forms2 but there also 
appear to be splay-footed groups of     
,pterodactyles like the species which   
have been named ,p4 elegans and ,p4     
micronyx1 in which the metatarsus widens
 out so that the bones of the toes do   
not diverge1 and that condition         

characterises the ,ptenodracon          
7,pterodactylus brevirostris71 to which 
genus these species may possibly be     
referred4 ,nearly all who have studied  
these animals regard the singularly     
short-nosed species ,p4 brevirostris as 
forming a separate genus4 ,for that     
genus ,soemmerring's descriptive name   
,ornithocephalus1 which he used for     
,pterodactyles generally1 might perhaps 
have been retained4 ,but the name       
,ptenodracon1 suggested by ,mr4         
,lydekker1 has been used for these      
types4    ,illustration3 ,,fig4 #fb4    
,,cycnorhamphus ,,suevicus    ,skeleton 
restored from the bones in ,fig4 #fj    
,illustration3 ,,fig4 #fc4              
,,restoration ,,of ,,skeleton           
,,cycnorhamphus ,,fraasi   ,,showing    
,,the ,,limbs ,,on ,,the ,,right ,,side 
,from a specimen in the ,museum at      
,stuttgart    ,illustration3 ,,fig4 #fd4
  ,,cycnorhamphus ,,fraasi              
,,restoration ,,of ,,the ,,form ,,of    
,,the ,,body  ,some of the largest      

specimens preserved at ,stuttgart and   
,tuebingen have been named              
,pterodactylus suevicus and ,p4         
,fraasii4 ,they do not approach the     
species ,p4 grandis in size1 so far as  
can be judged from the fragmentary      
remains figured by ,von ,meyer2 for what
 appears to be the third phalange of the
 wing finger is #g-a_/b inches long1    
while in these species it is less than  
half that length1 indicating an enormous
 development of wing1 relatively to the 
length of the hind limb1 which would    
probably refer the species to another   
genus4 ,pterodactylus suevicus differs  
from the typical ,pterodactyles in      
having a rounded1 flattened under       
surface to the lower jaw1 instead of the
 common condition of a sharp keel in the
 region of the symphysis4 ,the beak also
 seems flattened and swan-like1 and the 
teeth are limited to the front of the   
jaw4 ,there appear to be some           
indications of small nostrils1 which    
look upward like the nostrils of        

,rhamphorhynchus1 but this may be a     
deceptive appearance1 and the nostrils  
are large lateral vacuities1 which are  
in the position of antorbital vacuities1
 so that there would appear to be only  
two vacuities in the side of the head in
 these animals4 ,the distinctive        
character of the skeleton in this genus 
is found in the extraordinary length of 
the metacarpus and in the complete      
ossification of the smaller metacarpal  
bones throughout their length4 ,the     
metacarpal bones are much longer than   
the bones of the fore-arm1 and about    
twice the length of the humerus4 ,the   
first wing phalange is much longer than 
the others1 which successively and      
rapidly diminish in length1 so that the 
third is half the length of the first4  
,there are differences in the pelvis2   
for the anterior process of the ilium is
 very short1 in comparison with its     
length in the genus ,pterodactylus4 ,and
 the long stalk of the prepubic bone    
with its great hammer-headed expansion  

transversely in front gives those bones 
a character unlike other genera1 so that
 ,cycnorhamphus ranks as a good genus1  
easily distinguished from ,cuvier's     
type1 in which the four bones of the    
wing are more equal in length1 and the  
last is more than half the length of the
 first2 while the metacarpus in that    
genus is only a little longer than the  
humerus1 and much shorter than the ulna4
 ,the ,pterodactylus suevicus has the   
neck vertebrae flat on the under side1  
and relatively short as compared with   
the more slender and narrower vertebrae 
of ,p4 ,fraasii4     ,,chapter ,,xv     
,,ornithosaurs ,,from ,,the ,,upper     
,,secondary ,,rocks   ,when staying at  
,swanage1 in ,dorsetshire1 many years   
ago1 ,i had the rare good fortune to    
obtain from the ,purbeck ,beds the jaw  
of a ,pterodactyle1 which had much in   
common in plan with the ,cycnorhamphus  
,fraasii from the ,lithographic ,slate1 
which is preserved at ,stuttgart4 ,the  
tooth-bearing part of this lower jaw is 

#h inches long as preserved1 extending  
back #c inches beyond the symphysis     
portion in which the two sides are      
blended together4 ,it is different from 
,professor ,fraas's specimen in having  
the teeth carried much further back1 and
 in the animal being nearly twice as    
large4 ,this fragment of the jaw is     
little more than #a foot long1 which is 
probably less than half its original    
length4 ,a vertebra nearly #e inches    
long1 which is more than twice the      
length of the longest neck bones in the 
,stuttgart fossil1 is the only          
indication of the vertebral column4     
,professor ,owen described a wing finger
 bone from these ,purbeck ,beds1 which  
is nearly #a foot long4 ,he terms it the
 second of the finger4 ,it may be the   
third1 and on the hypothesis that the   
animal had the proportions of the       
,solenhofen fossil just referred to1 the
 first wing finger bone of the ,english 
,purbeck ,pterodactyle would have       
exceeded #b feet in length1 and would   

give a length for the wing finger of    
about #e feet #c inches4 ,for this      
animal the name ,doratorhynchus was     
suggested1 but at present ,i am unable  
to distinguish it satisfactorily from   
,cycnorhamphus1 which it resembles in   
the forms both of the neck bones and of 
the jaw4 ,very small ,pterodactyles are 
also found in the ,english ,purbeck     
strata1 but the remains are few1 and    
scattered1 like these larger bones4     
,illustration3 ,,fig4 #fe4  ,,the       
,,longest ,,known ,,neck ,,vertebra     
,from the ,purbeck ,beds of ,swanage4   
7,half natural size7   ,,ornithodesmus  
,,latidens    ,illustration3 ,,fig4 #ff4
  ,,cervical ,,vertebra ,,of            
,,ornithodesmus    ,from the ,wealden   
,beds of the ,isle of ,wight  ,the      
,wealden strata being shallow1          
fresh-water deposits might have been    
expected to supply better knowledge of  
,pterodactyles than has hitherto been   
available4 ,jaws of ,ornithocheirus     
sagittirostris have been found in the   

beds at ,hastings1 and in other parts of
 ,sussex4 ,some fragments are as large  
as anything known4 ,the best-preserved  
remains have come from the ,isle of     
,wight1 and were rewards to the         
enthusiastic search of the ,rev4 ,w4    
,fox1 of ,brixton4 ,in the principal    
specimen the teeth were short and wide1 
the head large and deep with large      
vacuities1 but the small brain case of  
that skull is bird-like4 ,the neck bones
 are #b-a_/b inches long4 ,in the upper 
part of the back the bones are united   
together by anchylosis1 so that they    
form a structure in the back like a     
sacrum1 which does not give attachment  
to the scapula1 as in some              
,pterodactyles from the ,chalk1 but the 
bones are simply blended1 as in the     
frigate-bird1 allied to ,pelicans and   
,cormorants4 ,and then after a few free 
vertebrae in the lower part of the back1
 succeeds the long sacrum1 formed in the
 usual way1 of many vertebrae4 ,i       
described a sacrum of this type from the

 ,wealden ,beds1 under the name         
,ornithodesmus1 referable to another    
species1 which in many respects was so  
like the sacrum of a ,bird that ,i could
 not at the time separate it from the   
bird type4 ,this genus has a sternum    
with a strong deep keel1 and the        
articulation for the coracoid bones     
placed at the back of the keel in the   
usual way1 but with a relation to each  
other seen in no genus hitherto known1  
for the articular surfaces are          
wedge-shaped instead of being ovate2 and
 instead of being side by side1 they    
obliquely overlap1 practically as in    
wading birds like the ,heron4 ,i have   
never seen any ,pterodactyle teeth so   
flattened and shaped like the end of a  
lancet2 and from this character the form
 was known between ,mr4 ,fox and his    
friends as latidens4 ,the name          
,ornithodesmus is as descriptive of the 
sternum as of the vertebral column4 ,the
 wing bones1 as far as they are         
preserved1 have the relatively great    

strength in the fore limb which is found
 in many of the ,pterodactyles of the   
,cretaceous period1 and are quite as    
large as the largest from the ,cambridge
 ,greensand4 ,in the ,sussex species    
named ,p4 sagittirostris the lower jaw  
articulation was inches wide4           
,illustration3 ,,fig4 #fg4  ,,sternum   
,,of ,,ornithodesmus    ,showing the    
overlapping facets for the coracoid     
bones 7shaded7 behind   the median keel 
,illustration3 ,,fig4 #fh4  ,,front ,,of
 ,,the ,,keel ,,of ,,the ,,sternum ,,of 
,,ornithodesmus ,,latidens    ,showing  
also the articulation for the coracoid  
bone  ,a few ,pterodactyles' bones have 
been discovered in the ,neocomian sands 
of ,england and ,germany1 and other     
larger bones occur in the ,gault of     
,folkestone and the north of ,france2   
but never in such association as to     
throw light on the aspect of the        
skeleton4   ,,ornithocheirus  ,within my
 own memory ,pterodactyle remains were  
equally rare from the ,cambridge        

,greensand4 ,the late ,professor ,owen  
in one of his public lectures produced  
the first few fragments received from   
,cambridge1 and with a knowledge which  
in its scientific method seemed to      
border on the power of creation1        
produced again the missing parts1 so    
that the bones told their story1 which  
the work of waves and mineral changes in
 the rock had partly obliterated4       
,subsequently good fortune gave me the  
opportunity during ten years to help my 
,university in the acquisition and      
arrangement of the finest collection of 
remains of these animals in ,europe4    
,out of an area of a few acres1 during a
 year or two1 came the thousand bones of
 ,ornithosaurs1 mostly associated sets  
of remains1 each a part of a separate   
skeleton1 described in my published     
catalogues1 as well as the best of those
 at ,york and in the ,british ,museum   
and other collections in ,london4  ,the 
deposit which yields them1 named        
,cambridge ,greensand1 may or may not   

represent a long period of time in its  
single foot of thickness2 but the       
abundance of fossils1 obtained whenever 
the workmen were adequately remunerated 
for preserving them1 would suggest that 
the ,pterodactyles might have lived like
 sea-birds or in colonies like the      
,penguins1 if it were not that the      
number of examples of each species found
 is always small1 and the many          
variations of structure suggested rather
 that the individuals represent the life
 of many lands4 ,the collections of     
remains are mostly from villages in the 
immediate vicinity of ,cambridge1 such  
as ,chesterton1 ,huntingdon ,road1      
,coldham ,common1 ,haslingfield1        
,barton1 ,shillington1 ,ditton1         
,granchester1 ,harston1 ,barrington1    
stretching south to ,ashwell in         
,bedfordshire on the one hand1 as well  
as further north by ,horningsea into the
 fens4 ,each appears to be the          
associated bones of a single individual4
 ,the remains mostly belong to          

comparatively large animals4 ,some were 
small1 though none have been found so   
diminutive as the smallest from the     
,solenhofen ,slate4 ,the largest        
specimens with long jaws appear to have 
had the head measuring not more than    
eighteen inches in length1 which is less
 than half the size of the great        
toothless ,pterodactyles from ,kansas4  
,illustration3 ,,fig4 #fi4              
,,restoration ,,of ,,the ,,skull ,,of   
,,ornithocheirus    ,the parts left     
white are in the ,geological ,museum at 
,cambridge4 ,the   shaded parts have not
 been found4 ,the two holes are the eye 
and the   nostril 7,from the ,cambridge 
,greensand7  ,the ,cambridge specimens  
manifestly belong to at least three     
genera4 ,something may be said of the   
characters of the large animals which   
are included in the genus               
,ornithocheirus4 ,these fossils have    
many points of structure in common with 
the great ,american toothless forms     
which are of similar geological age4    

,the skull is remarkable for having the 
back of the head prolonged in a         
compressed median crest1 which rose     
above the brain case1 and extended      
upward and over the neck vertebrae1 so  
as to indicate a muscular power not     
otherwise shown in the group4 ,for about
 three inches behind the brain this     
wedge of bone rested on the vertebrae1  
and probably overlapped the first three 
neural arches in the neck4  ,another    
feature of some interest is the         
expansion of the bone which comes below 
the eye4 ,in ,birds this malar or cheek 
bone is a slender rod1 but in these     
,pterodactyles it is a vertical plate1  
which is blended with the bone named the
 quadrate bone1 which makes the         
articulation with the lower jaw in all  
oviparous animals4  ,the beak varies    
greatly in length and in form1 though it
 is never quite so pointed as in the    
,american genus1 for there is always a  
little truncation in front1 when teeth  
are seen projecting forward from a      

position somewhat above the palate2 the 
snout is often massive and sometimes    
club-shaped4 ,except for these          
variations of shape in the compressed   
snout1 which is characterised by a ridge
 in the middle of the palate1 and a     
corresponding groove in the lower jaw1  
and the teeth1 there is little to       
distinguish what is known of the skull  
in its largest ,english ,greensand      
fossils from the skull remains which    
abound in the ,chalk of ,kansas4  ,this 
,english genus ,ornithocheirus1         
represented by a great number of        
species1 had the neural arch of the neck
 bones expanded transversely over the   
body of the vertebra in a way that      
characterises many birds with powerful  
necks1 and is seen in a few             
,pterodactyles from ,solenhofen4  ,it is
 difficult to resist the conclusion that
 the neck vertebrae were not usually    
more than twice to three times as long  
as those of the back1 and it would      
appear that the caudal vertebrae in the 

,english ,cretaceous types were         
comparatively large1 and about twice as 
long as the dorsal vertebrae4 ,unless   
there has been a singular succession of 
accidents in the association of these   
vertebrae with the other remains1       
,ornithocheirus had a tail of moderate  
length1 formed of a few vertebrae as    
long as those of the neck1 though more  
slender1 quite unlike the tail in either
 the long-tailed or short-tailed groups 
of ,solenhofen ,pterodactyles1 and      
longer than in the toothless            
,pterodactyles of ,america4             
,illustration3 ,,fig4 #gj4  ,,cervical  
,,vertebra1 ,,ornithocheirus    ,under  
side1 half natural size4 7,cambridge    
,greensand7  ,the singular articulation 
for the humerus at the truncated        
extremity of the coracoid bone is a     
character of this group1 as is the      
articulation of the scapulae with the   
neural arches of the dorsal vertebrae1  
at right angles to them 7p4 #aae71      
instead of running over the ribs as in  

,birds and as in other ,pterodactyles4  
,the smaller ,pterodactyles have their  
jaws less compressed from side to side4 
,the upper arm bone1 the humerus1       
instead of being truncated at its lower 
end as in ,ornithocheirus1 is divided   
into two or three rounded articular     
surfaces4 ,that for the radius1 the bone
 which carries the wrist1 is a distinct 
and oblique rounded facet1 while the    
ulna has a rounded and pulley-like      
articulation on which the hand may      
rotate4 ,these differences are probably 
associated with an absence of the       
remarkable mode of union of the scapulae
 with the dorsal vertebrae4 ,but ,i have
 hesitated to give different names to   
these smaller genera because no example 
of scapula has come under my notice     
which is not truncated at the free end4 
,i do not think this ,european type can 
be the ,nyctodactylus of ,professor     
,marsh1 in which sutures appear to be   
persistent between the bodies of the    
vertebrae and their arches1 because no  

examples have been found at ,cambridge  
with the neural arches separated1       
although the scapula is frequently      
separated from the coracoid in large    
animals4    ,illustration3 ,,fig4 #ga4  
,,upper ,,and ,,lower ,,jaws ,,of ,,an  
,,english   ,,pterodactyle ,,from ,,the 
,,chalk1 ,,as ,,preserved               
,,ornithostoma    ,illustration3 ,,fig4 
#gb4  ,,the ,,palate ,,of ,,the         
,,english ,,toothless   ,,pterodactyle1 
,,ornithostoma    ,illustration3 ,,fig4 
#gc4  ,,types ,,of ,,the ,,american     
,,toothless ,,pterodactyle1             
,,ornithostoma    ,named by ,marsh1     
,pteranodon  ,the most interesting of   
all the ,english ,pterodactyle remains  
is the small fragment of jaw figured by 
,sir ,richard ,owen in #ahei1 which is a
 little more than two inches long and an
 inch wide1 distinguished by a concave  
palate with smooth rounded margins to   
the jaws and a rounded ridge to the     
beak4 ,it is the only satisfactory      
fragment of the animal which has been   

figured1 and indicates a genus of       
toothless ,pterodactyles1 for which the 
name ,ornithostoma was first used in    
#ahga4 ,after some years ,professor     
,marsh found toothless ,pterodactyles in
 ,kansas1 and indicated several species4
 ,there are remains to the number of six
 hundred specimens of these ,american   
animals in the ,yale ,museum alone2 but 
very little was known of them till      
,professor ,williston1 of ,lawrence1 in 
,kansas1 described the specimens from   
the ,kansas ,university ,museum1 when it
 became evident that the bones of the   
skeleton are mostly formed on the same  
plan as those of the ,cambridge         
,greensand genus1 ,ornithocheirus4 ,they
 are not quite identical4 ,professor    
,williston adopts for them the name     
,ornithostoma1 in preference to         
,pteranodon which ,marsh had suggested4 
,both animals have the dagger-shaped    
form of jaw1 with corresponding height  
and breadth of the palate4 ,the same    
flattened sides to the snout1 converging

 upwards to a rounded ridge1 the same   
compressed rounded margin to the jaw1   
which represents the border in which    
teeth are usually implanted1 and in both
 the palate has the same smooth         
character forming a single wide concave 
channel4 ,years previously ,i had the   
pleasure of showing to ,professor ,marsh
 the remarkable characters of the jaw1  
shoulder-girdle bones1 and scapulae in  
the ,greensand ,pterodactyles while the 
,american fossils were still            
undiscovered4 ,i subsequently made the  
restoration of the shoulder-girdle 7p4  
#aae74 ,professor ,williston states to  
me that the shoulder-girdle bones in    
,american examples of ,ornithostoma have
 a close resemblance to those of        
,ornithocheirus figured in #ahia1 as is 
evident from remains now shown in the   
,british ,museum4 ,it appears that the  
,kansas bones are almost invariably     
crushed flat1 so that their articular   
ends are distorted4 ,the neck vertebrae 
are relatively stout as in              

,ornithocheirus4 ,the hip-girdle of the 
,american ,ornithostoma can be closely  
paralleled in some ,english specimens of
 ,ornithocheirus1 though each prepubic  
bone is triangular in the ,american     
fossils as in ,p4 rhamphastinus4 ,they  
are united into a transverse bar as in  
,rhamphorhynchus1 unknown in the        
,english fossils4 ,the femur has the    
same shape as in ,ornithocheirus2 and   
the long tibia terminates in a pulley4  
,there is no fibula4 ,the sternum in    
both has a manubrium1 or thick keel     
mass1 prolonged in front of its         
articular facets for the coracoid bones1
 which are well separated from each     
other4 ,four ribs articulate with its   
straight sides4 ,the animal has four    
toes and the fifth is rudimentary2 there
 are no claws to the first and second4  
,illustration3 ,,fig4 #gd4              
,,restoration ,,of ,,the ,,skeleton ,,of
 ,,ornithostoma   ,,ingens 7,,marsh7    
,from the ,niobrara ,cretaceous of      
,western ,kansas4 ,made by ,professor   

,williston4 ,the original has a spread  
of wing of about #ai feet #d   inches4  
,fragments of larger individuals are    
preserved at ,munich  ,in the           
restoration which ,professor ,williston 
has made the wing metacarpal is long1   
and in the shortest specimen measures #a
 foot #g inches1 and in the longest #a  
foot #h inches4 ,this is exactly equal  
to the length of the first phalange of  
the wing finger4 ,the second wing finger
 bone is #c inches shorter1 the third is
 little more than half the length of the
 first1 while the fourth is only #f-c_/d
 inches long1 showing a rapid shortening
 of the bones1 a condition which may    
have characterised all the ,cretaceous  
,pterodactyles4 ,the short humerus1     
about #a foot long1 and the fore-arm1   
which is scarcely longer1 are also      
characteristic proportions of           
,ornithostoma or ,pteranodon1 as known  
from the ,american specimens4 ,professor
 ,williston gives no details of the     
remarkable tail1 beyond saying that the 

tail is small and short1 and that the   
vertebrae are flat at the ends1 without 
transverse processes4 ,in the           
restoration the tail is shorter than in 
the short-tailed species from the       
,lithographic ,slate1 and unlike the    
tail in ,ornithocheirus4   ,this is the 
succession of ,pterodactyles in         
geological time4 ,their history is like 
that of the human race4 ,in the most    
ancient nations man's life comes upon us
 already fully organised4 ,the          
,pterodactyles begin1 so far as isolated
 bones are concerned1 in the ,rhaetic   
strata2 perhaps in the ,muschelkalk or  
middle division of the ,trias4 ,and from
 the beginning of the ,secondary time   
they live on with but little diversity  
in important and characteristic         
structures1 and so far as habit goes1   
the great ,pterodactyles of the ,upper  
,chalk of ,england cannot be said to be 
more highly organised than the earlier  
stiff-tailed genera of the ,lias or the 
,oolites4 ,there is nothing like        

evolution4 ,no modification such as that
 which derives the one-toed horse or the
 two-toed ox from ancestors with a      
larger number of digits4 ,on the other  
hand1 there is little1 if any1 evidence 
of degeneration4 ,the later             
,pterodactyles do not appear to have    
lost much1 although the tail in some of 
the ,solenhofen genera may be degenerate
 when compared with the long tail of    
,dimorphodon2 but the short-tailed types
 are found side by side with the        
long-tailed ,rhamphorhynchus4 ,the      
absence of teeth may be regarded as     
degeneration1 for they have presumably  
become lost1 in the same way that ,birds
 now existing have lost the teeth which 
characterised the fossil                
birds--,ichthyornis of the ,american    
,greensand1 and ,archaeopteryx of the   
,upper ,oolites of ,bavaria4 ,but just  
as some of the earlier ,pterodactyles   
have no teeth at the extremity of the   
jaw1 such as ,dorygnathus and           
,rhamphorhynchus1 so the loss of teeth  

may have extended backward till the jaws
 became toothless4 ,the specimens       
hitherto known give no evidence of such 
a change being in progress4 ,but just as
 the division of ,mammals termed        
,edentata usually wants only the teeth  
which characterise the front of the jaw1
 yet others1 like the ,great ,ant-eater 
of ,south ,america named ,myrmecophaga1 
have the jaws as free from teeth as the 
toothless ,pterodactyles or living      
,birds1 and show that in that order the 
teeth have no value in separating these 
animals into subordinate groups any more
 than they have among the ,monotremata1 
where one type has teeth and the other  
is toothless4  ,the following table     
gives a summary of the ,geological      
,history and succession in the          
,secondary ,rocks of the principal      
genera of ,flying ,reptiles4            
----------------------------------------
-----------------------------           
,,names ,,of ,,the ,,genera4            
,,geological ,,formations4              

----------------------------------------
-----                           ,british
 and ,european4  ,north ,american4      
----------------------------------------
----------------------------   ,upper   
,chalk                                  
,ornithostoma                           
7,pteranodon7   ,lower ,chalk           
,ornithocheirus        ,nyctodactylus   
,upper ,greensand                       
,ornithostoma       ,gault              
-----------------------                 
,lower ,greensand                       
,wealden                 ,ornithodesmus 
,purbeck                 ,doratorhynchus
           -----------------------      
,portland                ,pterodactylus 
,ptenodracon             ,kimeridge     
,clay and      ,cycnorhamphus           
,solenhofen ,slate      ,diopecephalus  
,rhamphorhynchus         ,coralline     
,oolite        ,scaphognathus           
,oxford ,clay                           
-----------------------                 
,great ,oolite and                      

,stonesfield ,slate     ,rhamphocephalus
                                        
,inferior ,oolite                       
-----------------------                 
,upper ,lias                            
,campylognathus                         
,dorygnathus             ,lower ,lias   
,dimorphodon                            
-----------------------                 
,rhaetic                   bones        
,muschelkalk              8 bones       
----------------------------------------
----------------------------            
,,chapter ,,xvi  ,,classification ,,of  
,,the ,,ornithosauria   ,when an attempt
 is made to determine the place in      
nature of an extinct group of animals   
and the relation to each other of the   
different types included within its     
limits1 so as to express those facts in 
a classification1 attention is directed 
in the first place to characters which  
are constant1 and persist through the   
whole of its constituent genera4 ,we    
endeavour to find the structural parts  

of the skeleton which are not affected  
by variation in the dentition1 or the   
proportions of the extremities1 or      
length of the tail1 which may define    
families or genera1 or species4  ,it has
 already been shown that while in many  
ways the ,ornithosaurian animals are    
like ,birds1 they have also important   
resemblances to ,reptiles4 ,they are    
often named ,pterosauria4 ,the wing     
finger gives a distinctive character    
which is found in neither one class of  
existing animals nor the other1 and is  
common to all the ,pterodactyles at     
present known4 ,they have been named    
,ornithosauria as a distinct minor      
division of back-boned animals1 which   
may be regarded as neither ,reptiles nor
 ,birds in the sense in which those     
terms are used to define a ,lizard or   
,ostrich among animals which still      
exist4 ,it is not so much that they mark
 a transition from ,reptile to ,bird1 as
 that they are a group which is parallel
 to ,birds1 and more manifestly holds an

 intermediate place than ,birds do      
between ,reptiles and ,mammals4 ,in plan
 of structure ,bird and ,reptile have   
more in common than was at one time     
suspected4 ,the late ,professor ,huxley 
went so far as to generalise on those   
coincidences in parts of the skeleton1  
and united ,birds and ,reptiles into one
 group1 which he named ,sauropsida1 to  
express the coincidences of structure   
between the ,lizard and the ,bird       
tribes4 ,the idea is of more value than 
the term in which it is expressed1      
because ,reptiles are not1 as we have   
seen1 a group of animals which can be   
defined by any set of characters as     
comprehensive as those which express the
 distinctive features of ,birds4 ,from  
the anatomist's point of view ,birds are
 a smaller group1 and while some        
,reptiles have affinity with them1 it is
 rather the extinct than the living     
groups which indicate that relation4    
,other ,reptiles have affinities of a   
more marked kind with ,mammals1 and     

there are points in the ,ornithosaurian 
skeleton which are distinctly           
,mammalian4 ,so that when the ,monotreme
 ,mammals are united with ,south        
,african reptiles known as              
,theriodontia1 which resemble them1 in a
 group termed ,theropsida to express    
their mammalian resemblances1 it is     
evident that there is no one continuous 
chain of life or gradation in complexity
 of structure of animals4  ,we have to  
determine whether the ,ornithosauria    
incline towards the ,sauropsidan or     
,bird-,reptile alliance1 or to the      
,mammal-,reptile or ,theropsidan        
alliance4 ,there can be no doubt that   
the predominant tendency is to the      
former1 with a minor affinity towards   
the latter4  ,the ,ornithosauria are one
 of a series of groups of animals1      
living and extinct1 which have been     
combined in an alliance named the       
,ornithomorpha4 ,that group includes at 
least five great divisions of animals1  
which circle about birds1 known as      

,ornithosauria1 ,crocodilia1            
,saurischia1 ,aves1 ,ornithischia1 and  
,aristosuchia4 ,their relations to each 
other are not evident in an enumeration1
 but may be shown in some degree in a   
diagram 7see p4 #aij74   ,,the          
,,ornithomorpha  ,the ,ornithomorpha    
arranged in this way show that the three
 middle groups--carnivorous ,saurischia1
 ,aristosuchia1 herbivorous             
,ornithischia--which are usually united 
as ,dinosauria1 intervene between ,birds
 and ,ornithosaurs2 and that the        
,crocodilia and ,ornithosauria are      
parallel groups which are connected with
 ,birds1 by the group of ,dinosaurs1    
which resembles ,birds most closely4    
,the ,ornithomorpha is only one of a    
series of large natural groups of       
animals into which living and extinct   
terrestrial vertebrata may be arranged4 
,and the succeeding diagram may         
contribute to make evident the relations
 of ,ornithosauria to the other         
terrestrial vertebrata 7see p4 #aia74   

,herein it is seen that while the       
,ornithomorpha approach towards         
,mammalia through the ,ornithosauria1   
and less distinctly through the         
,crocodilia1 they approach more directly
 to the ,sauromorpha1 through the       
,plesiosaurs and ,hatteria2 while that  
group also approaches more directly to  
the ,mammals through the ,plesiosaurs   
and ,anomodonts4    ,illustration3      
,,diagram ,,of ,,the ,,affinities ,,of  
,,the ,,orders ,,of ,,animals           
,,comprised ,,in ,,the ,,ornithomorpha4 
,after a diagram in the ,philosophical  
,transactions of the ,royal   ,society1 
#ahib4  ,the ,aristosuchia is           
imperfectly known1 and therefore to some
 extent a provisional group4 ,it is a   
small group of animals4                 
,illustration3 ,,diagram ,,showing ,,the
 ,,relations ,,of ,,the ,,ornithomorpha 
,,to ,,the ,,chief ,,large ,,groups ,,of
 ,,terrestrial ,,vertebrata1 ,,and      
,,their   ,,affinities ,,with ,,each    
,,other4    ,after a diagram in the     

,philosophical ,transactions of the     
,royal   ,society1 #ahib4               
,cordylomorpha are ,ichthyosaurs and the
 ,labyrinthodont group4 ,herpetomorpha  
include ,lacertilia1 ,homoeosauria1     
,dolichosauria1 ,chameleonoidea1        
,ophidia1 ,pythonomorpha4  ,the         
,sauromorpha comprises the groups of    
extinct and living ,reptiles named      
,chelonia1 ,rhynchocephala1             
,sauropterygia1 ,anomodontia1           
,nothosauria1 and ,protorosauria4 ,these
 details may help to explain the place  
which has been given to the             
,ornithosauria in the classification of 
animals4    ,illustration3 ,,fig4 #ge4  
,,comparison ,,of ,,six ,,genera    ,the
 skulls are seen on the left side in the
 order of the names below   them        
,turning to the ,pterodactyles          
themselves1 ,von ,meyer divided them    
naturally into short-tailed and         
long-tailed4 ,the short-tailed indicated
 by the name ,pterodactylus he further  
divided into long-nosed and short-nosed4

 ,the short-nosed genus has since been  
named ,ptenodracon 7,fig4 #ei1 p4 #afg74
 ,the long-tailed group was divided into
 two types--the ,rhamphorhynchus of the 
,solenhofen ,slate 7,fig4 #ef1 p4 #afa7 
and the ,english form now known as      
,dimorphodon 7,fig4 #eb1 p4 #aej71 which
 had been described from the ,lias4     
,the ,cretaceous ,pterodactyles form a  
distinct family4 ,so that1 believing the
 tail to have been short in that group  
7,fig4 #eh71 there are two long-tailed  
as well as two short-tailed families1   
which were defined from their typical   
genera ,pterodactylus1 ,ornithocheirus1 
,rhamphorhynchus1 and ,dimorphodon4     
,the differences in structure which     
these animals present are1 first3 the   
big-headed forms from the ,lias like    
,dimorphodon1 agree with the            
,rhamphorhynchus type from ,solenhofen  
in having a vacuity in the skull defined
 by bone1 placed between the orbit of   
the eye and the nostril4 ,with those    
characters are correlated the           

comparatively short bones which         
correspond to the back of the hand      
termed metacarpals1 and the tail is     
long1 and stiffened down its length with
 ossified tendons4 ,these characters    
separate ,ornithosaurs with long tails  
from those with short tails4  ,the      
short-tailed types represented by       
,pterodactylus and ,ornithocheirus have 
no distinct antorbital vacuity in the   
skull defined by bone4 ,the metacarpal  
bones of the middle hand are            
exceptionally elongated1 and the tail1  
which was flexible in both1 appears to  
have been short4 ,these differences in  
the skeleton warrant a primary division 
of flying reptiles into two principal   
groups4  ,the short-tailed group1 which 
was recognised by ,de ,blainville as    
intermediate between ,birds and         
,reptiles1 may take the name            
,pterodactylia1 which he suggested as a 
convenient1 distinctive name4 ,it may   
probably be inconvenient to enlarge its 
significance to comprise not only the   

true ,pterodactyles originally defined  
as ,pterosauria1 but the newer          
,ornithostoma and ,ornithocheirus which 
have been grouped as ,ornithocheiroidea4
  ,the second order1 in which the wing  
membrane appears to have had a much     
greater extent1 in being carried down   
the hind limbs1 where the outermost     
digit and metatarsal are modified for   
its support1 has been named             
,pterodermata1 to include the types     
which are arranged around               
,rhamphorhynchus and ,dimorphodon4      
,both these principal groups admit of   
subdivision by many characters in the   
skeleton1 the most remarkable of which  
is afforded by the pair of bones carried
 in front of the pubes1 and termed      
prepubic bones4 ,in the ,pterodactyle   
family the bones in front of the pubes  
are always separate from each other1    
always directed forward1 and have a     
peculiar fan-shaped form with concave   
sides like the bone which holds a       
similar position in a ,crocodile4 ,in   

the ,ornithocheirus family the prepubic 
bones appear to have been originally    
triangular1 but were afterwards united  
so as to form a strong continuous bar   
which extends transversely across the   
abdomen in advance of the pubic bones4  
,this at least is the distinctive       
character in the genus ,ornithostoma    
according to ,professor ,williston1     
which in many ways closely resembles    
,ornithocheirus4  ,the two families in  
the long-tailed order named             
,pterodermata are separated from each   
other by a similar difference in their  
prepubic bones4 ,in ,dimorphodon those  
bones are separate from each other1 and 
remain distinct through life1 meeting in
 the middle line of the body in a wide  
plate4 ,on the other hand1 in           
,rhamphorhynchus the prepubic bones1    
which are at first triangular and always
 slender1 become blended together into a
 slight transverse bar1 which only      
differs from that attributed to         
,ornithostoma in its more slender       

bow-shaped form4    ,illustration3      
,,fig4 #gf4  ,,left ,,side ,,of ,,pelvis
 ,,of ,,ornithostoma   7,after          
,williston7  ,thus if other characters  
of the skeleton are ignored and a       
classification based upon the structure 
of the pelvis and prepubic bones1 there 
would be some ground for associating the
 long-tailed ,rhamphorhynchus from the  
,upper ,oolites which is losing the     
teeth in the front of its jaw with the  
,cretaceous ,ornithostoma1 which has the
 teeth completely wanting2 while the    
long-tailed ,dimorphodon would come into
 closer association with the            
short-tailed ,pterodactylus4 ,the       
drum-stick bone or tibia in             
,dimorphodon1 with its slender fibula1  
like that of a ,bird1 also resembles a  
,bird in the rounded and pulley-shaped  
terminal end which makes the joint      
corresponding to the middle of the ankle
 bones in man4 ,the same condition of a 
terminal pulley joint is found in the   
,cretaceous ,pterodactyles4 ,but in the 

true ,pterodactyles and in              
,rhamphorhynchus there usually is no    
pulley-shaped termination to the lower  
end of the drum-stick1 for the tarsal   
bones remain separate from each other1  
and form two rows of ossifications1     
showing the same differences as separate
 ,dinosaurs into the divisions which    
have been referred to1 from their       
,bird-like pelvis and tibio-tarsus1 as  
,ornithischia in the one case1 and      
,saurischia in the other from their     
bones being more like those of living   
,lizards4     ,,chapter ,,xvii  ,,family
 ,,relations ,,of ,,pterodactyles ,,to  
,,animals ,,which ,,lived ,,with ,,them 
,enough has been said of the general    
structure of ,pterodactyles and the     
chief forms which they assumed while the
 ,secondary rocks were accumulating1 to 
convey a clear idea of their relations  
to the types of vertebrate animals which
 still survive on the earth4 ,we may be 
unable to explain the reasons for their 
existence1 and for their departure from 

the plan of organisation of ,reptiles   
and ,birds4 ,but the evidence has not   
been exhausted which may elucidate their
 existence4 ,sometimes1 in problems of  
this kind1 which involve comparison of  
the details of the skeleton in different
 animals1 it is convenient to imagine   
the possibility of changes and          
transitions which are not yet supported 
by the discovery of fossil remains4 ,if1
 for example1 the ,pterodactyle be      
conceived of as divested of the wing    
finger1 which is its most distinctive   
character1 or that finger is supposed to
 be replaced by an ordinary digit1 like 
the three-clawed digits of the hand     
which we have regarded as applied to the
 ground1 where1 it may be asked1 would  
the animal type be found which          
approximates most closely to a          
,pterodactyle which had been thus       
modified8 ,there are two possible       
replies to such a question1 suggested by
 the form of the foot4 ,for the old     
,bird ,archaeopteryx has three such     

clawed digits1 but no wing finger4 ,and 
some ,dinosaurs also have the hand with 
three digits terminating in claws1 which
 are quite comparable to the clawed     
digits of ,pterodactyles4  ,the truth   
expressed in the saying that no man by  
taking thought can add a cubit to his   
stature is of universal application in  
the animal world1 in relation to the    
result upon the skeleton of the exercise
 of a function by the individual4 ,yet  
such is the relation in proportions of  
the different parts of the animal to the
 work which it performs1 so marked is   
the evidence that growth has extended in
 direct relation to use of organs and   
active life1 and that structures have   
become dwarfed from overwork1 or have   
wasted away from disuse--seen throughout
 all vertebrate animals1 that we may    
fairly attribute to the wing finger some
 correlated influence upon the          
proportions of the animal1 as a         
consequence of the dependence of the    
entire economy upon each of its parts4  

,therefore if an allied animal did not  
possess a wing finger1 and did not fly1 
it might not have developed the         
lightness of bone1 or the length of limb
 which ,pterodactyles possess4  ,the    
mere expansion of the parachute membrane
 seen in so-called flying animals1 both 
,mammals and ,reptiles1 which are devoid
 of wings1 is absolutely without effect 
in modifying the skeleton4 ,but when in 
the ,bat a wing structure is met with   
which may be compared to a gigantic     
extension of the web foot of the        
so-called ,flying ,frog1 the bones of   
the fingers and the back of the hand    
elongate and extend under the stimulus  
of the function of flight in the same   
way as the legs elongate in the more    
active hoofed animals1 with the function
 of running4 ,therefore it is not       
improbable that the limbs shared to some
 extent in growth under stimulus of     
exercise which developed the wing       
finger4 ,and if an animal can be found  
among fossils so far allied as to       

indicate a possible representative of   
the race from which these ,flying       
,dragons arose1 it might be expected to 
be at least shorter legged1 and possibly
 more distinctly ,reptilian in the bones
 of the shoulder-girdle which support   
the muscles used in flight4 ,it may     
readily be understood that the kinds of 
life which were most nearly allied to   
,pterodactyles are likely to have       
existed upon the earth with them1 and   
that flight was only one of the modes of
 progression which became developed in  
relation to their conditions of         
existence4 ,the principal assemblage of 
terrestrial animals available for such  
comparison is the ,dinosauria4 ,they may
 differ from ,pterodactyles as widely as
 the ,insectivora among ,mammals differ 
from ,bats1 but not in a more marked    
way4 ,comparisons will show that there  
are resemblances between the two extinct
 groups which appeal to both reason and 
imagination4  ,dinosaurs are            
conveniently divided by characters of   

the pelvis first into the order         
,saurischia1 which includes the         
carnivorous ,megalosaurus and the       
,cetiosaurus1 with the pelvis on the    
,reptile plan2 and secondly the order   
,ornithischia1 represented by           
,iguanodon1 with the pelvis on the ,bird
 plan4 ,it may be only a coincidence1   
but nevertheless an interesting one1    
that the characters of those two great  
groups of reptiles1 which also extend   
throughout the ,secondary rocks1 are to 
some extent paralleled in parts of the  
skeleton of the two divisions of        
,pterodactyles4 ,this may be illustrated
 by reference to the skull1 pelvis1 hind
 limb1 and the pneumatic condition of   
the bones4    ,illustration3 ,,fig4 #gg4
  ,,comparison ,,of ,,the ,,skull ,,of  
,,the ,,dinosaur   ,,anchisaurus ,,with 
,,the ,,ornithosaur ,,dimorphodon  ,the 
,saurischian ,dinosauria have an        
antorbital vacuity in the side of the   
skull between the nasal opening and the 
eye1 as in the long-tailed ,ornithosaurs

 named ,pterodermata4 ,in some of the   
older genera of these carnivorous       
,dinosaurs of the ,trias1 the lateral   
vacuities of the head are as large as in
 ,dimorphodon4 ,but in some at least of 
the ,iguanodont1 or ,ornithischian      
,dinosaurs1 there is no antorbital      
vacuity1 and the side of the face in    
that respect resembles the short-tailed 
,pterodactylia4 ,the skull of a         
carnivorous ,dinosaur possesses teeth   
which1 though easily distinguished from 
those of ,pterodactyles1 can be best    
compared with them4 ,the most striking  
difference is in the fact that in the   
,dinosaur the nostrils are nearly       
terminal1 while in the ,pterodactyle    
they are removed some distance backward4
 ,this result is brought about by growth
 taking place1 in the one case at the   
front margin of the maxillary bone so as
 to carry the nostril forward1 and in   
the other case at the back margin of the
 premaxillary bone4 ,thus an elongated  
part of the jaw is extended in front of 

the nostril4 ,hence there is a different
 proportion between the premaxillary and
 maxillary bones in the two groups of   
animals1 which corresponds to the       
presence of a beak in a bird1 and its   
absence in living reptiles4 ,it is not  
known whether the extremity of the      
,pterodactyle's beak is a single bone1  
the intermaxillary bone1 such as forms  
the corresponding toothless part of the 
jaw in the ,south ,african reptile      
,dicynodon1 or whether it is made by the
 pair of bones called premaxillaries    
which form the extremity of the jaw in  
most ,dinosaurs4 ,too much importance   
may perhaps be attached to such         
differences which are partly            
hypothetical1 because the extinct       
,ichthyosaurus1 which has an            
exceptionally long snout1 has the two   
premaxillary bones elongated so as to   
extend backward to the nostrils4 ,a     
similar elongation of those bones is    
seen in ,porpoises1 which also have a   
long snout2 and the bones are carried   

back from the front of the head to the  
nostrils1 which are sometimes known as  
blowholes4 ,but the ,porpoise has those 
premaxillary bones not so much in       
advance of the bones which carry teeth  
named maxillary1 as placed in the       
interspace between them4 ,the nostrils1 
however1 are not limited to the         
extremity of the head in all ,dinosaurs4
 ,if this region of the beak in         
,dimorphodon be compared with the       
corresponding part of a ,dinosaur from  
the ,permian rocks1 or ,trias1 the      
relation of the nostril to the bones    
forming the beak may be better          
understood4    ,illustration3 ,,fig4    
#gh4  ,,comparison ,,of ,,the ,,skull   
,,of ,,the ,,dinosaur   ,,ornithosuchus 
,,with ,,the ,,ornithosaur ,,dimorphodon
  ,in the sandstone of ,elgin1 usually  
named ,trias1 a small ,dinosaur is      
found1 which has been named             
,ornithosuchus1 from the resemblance of 
its head to that of a ,bird4 ,seen from 
above1 the head has a remarkable        

resemblance to the condition in         
,rhamphorhynchus1 in the sharp-pointed  
beak and positions of the orbits and    
other openings4 ,in side view the orbits
 have the triangular form seen in       
,dimorphodon1 and the preorbital        
vacuities are large1 as in that genus1  
while the lateral nostrils1 which are   
smaller1 are further forward in the     
,dinosaur4 ,the differences from        
,dimorphodon are in the articulation for
 the jaw being carried a little         
backward1 instead of being vertical as  
in the ,pterodactyle1 and the bone in   
front of the nose is smaller4           
,notwithstanding probable differences in
 the palate1 the approximation1 which   
extends to the ,crocodile-like vacuity  
in the lower jaw1 is such that by slight
 modification in the skull the          
differences would be substantially      
obliterated by which the skull of such  
an ,ornithosaur is technically          
distinguished from such a ,dinosaur4    
,the back of the skull is clearly seen  

in the ,whitby ,pterodactyle1 and its   
structure is similar to the             
corresponding part of such ,dinosaurs as
 ,anchisaurus or ,atlantosaurus1 without
 the resemblance quite amounting to     
identity1 but still far closer than is  
the resemblance between the same region 
in the heads of ,crocodiles1 ,lizards1  
,serpents1 ,chelonians4 ,few of these   
fossil ,dinosaur skulls are available   
for comparison1 and those differ among  
themselves4 ,the coincidences rather    
suggest a close collateral relation than
 prove the elaboration of one type from 
the other4 ,they may have had a common  
ancestor4  ,the ,trias rocks near       
,stuttgart have yielded ,dinosaurs as   
unlike ,pterodactyles as could be       
imagined1 resembling heavily armoured   
,crocodiles1 in such types as the genus 
,belodon4 ,its jaws are compressed from 
side to side1 as in many ,pterodactyles1
 and the nostrils are at least as far   
backward as in ,rhamphorhynchus4        
,belodon has preorbital vacuities and   

postorbital vacuities1 but the orbit of 
the eye is never large1 as in           
,pterodactyles4 ,it might not be worth  
while dwelling on such points in the    
skull if it were not that the pelvis in 
,belodon is a basin formed by the       
blending of the expanded plates of the  
ischium and the pubis1 into a sheet of  
bone which more nearly resembles the    
same region in ,pterodactyles than does 
the ischio-pubic region in other        
,dinosaurian animals like ,cetiosaurus4 
,the backbone in a few ,dinosaurs is    
suggestive of ,pterodactyles4 ,in such  
genera as have been named ,coelurus and 
,calamospondylus1 in which the skeleton 
is only partially known1 the neck       
vertebrae become elongated1 so as to    
compare with the long-necked            
,pterodactyles4 ,the cervical rib is    
often very similar to that type1 and    
blended with the vertebra1 as in        
,pterodactyles and ,birds4 ,the early   
dorsal vertebrae of ,pterodactyles might
 almost be mistaken for those of        

,dinosaurs4 ,the tail vertebrae of a    
,pterodactyle are usually longer than in
 long-tailed ,dinosauria4  ,in the limbs
 and the bony girdles which support them
 there is more resemblance between      
,pterodactyles and ,dinosaurs than might
 have been anticipated1 considering     
their manifest differences in habit4    
,thus all ,dinosaurs have the hip bone  
named ilium prolonged in front of the   
articulation for the femur as well as   
behind it1 almost exactly as in         
,pterodactyles and ,birds 7see p4 #ie74 
,there is some difference in the pubis  
and ischium which is more conspicuous in
 form than in direction of the bones4   
,there is a ,pterodactyle imperfectly   
preserved1 named ,pterodactylus dubius1 
in which the ischium is directed        
backward and the pubis downward1 and the
 bones unite below the acetabular cavity
 for the head of the femur to work in1  
but do not appear to be otherwise       
connected4 ,in ,rhamphorhynchus the     
connexion between these two thickened   

bars is made by a thin plate of bone4   
,in such a ,dinosaur as the ,american   
carnivorous ,ceratosaurus the two bars  
of the pubis and ischium remain separate
 and diverging1 and there is no film of 
bone extending over the interspace      
between them4 ,the development of such a
 bony condition would make a close      
approximation between the               
,ornithosaurian pelvis and that of those
 ,dinosaurs which closely resemble      
,pterodactyles in skull and teeth4      
,illustration3 ,,fig4 #gi4  ,,left      
,,side ,,of ,,pelvis   ,a ,pterodactyle 
is shown between a carnivorous ,dinosaur
 above and a   herbivorous ,dinosaur    
below  ,another pelvic character of some
 interest is the blending of the pubis  
and ischium of the right and left sides 
in the middle line of the body4 ,there  
are some genera of ,dinosaurs like the  
,english ,aristosuchus from the ,weald1 
and the ,american genera ,coelurus1     
,ceratosaurus1 and others1 in which the 
pubic bones1 instead of uniting at their

 extremities1 are pinched together from 
side to side1 and unite down the lower  
part of their length1 terminating in an 
expanded end like a shoe1 which is seen 
to be a separate ossification1 and      
probably formed by a pair of            
ossifications joined in the median line4
 ,this small bone1 which is below the   
pubes1 and in these animals becomes     
blended with them1 we may regard as a   
pair of prepubic bones like those of    
,pterodactyles and ,crocodiles1 except  
that they have lost the stalk-like      
portions1 which in those animals are    
developed to compensate for the         
diminished length of the pubic bones4   
,the prepubic bones may also be         
developed in ,iguanodon1 in which a pair
 of bones of similar form remains       
throughout life in advance of the pubes1
 as in ,pterodactyles4 ,in those        
,dinosauria with the ,bird-like type of 
pelvis the pubic bone is exceptionally  
developed1 sending one process backward 
and another process forward1 so that    

there is a great gap between these      
diverging limbs to the bone4 ,in the    
region behind the sternum to which the  
ribs were attached1 and in front of the 
pelvis1 is a pair of bones in ,iguanodon
 shaped like the prepubic bones of      
,dimorphodon4 ,they have sometimes been 
interpreted as a hinder part of the     
sternum1 but may more probably be       
regarded as a pair of prepubic bones    
articulating each with the anterior     
process of the pubis 7see ,fig4 #hj74   
,the small bones found at the           
extremities of the pubes in such        
carnivorous ,dinosaurs as ,aristosuchus 
are blended by bony union with the      
pubes4 ,the bones in ,iguanodon are     
placed behind the sternal region without
 any attachment for sternal ribs1 and   
the expanded processes converge forwards
 from the stalk and unite exactly like  
the prepubic bones of ,ornithosaurs4    
,while this character1 on the one hand1 
may link ,pterodactyles with the        
,dinosaurs1 on the other hand it may be 

a link between both those groups and the
 ,crocodiles1 in which the front pair of
 bones of the pelvis has also appeared  
to be representative of the prepubic    
bones of ,flying ,reptiles 7see ,fig4   
#cb1 p4 #ih74    ,illustration3 ,,fig4  
#hj4  ,,diagram ,,of ,,the ,,pelvis     
,,seen ,,from ,,below ,,in ,,an         
,,ornithosaur ,,and ,a ,,dinosaur  ,the 
resemblances between ,pterodactyles and 
,dinosaurs in the hind limb are not of  
less interest1 though it is rather in   
the older ,pterodactyles such as        
,dimorphodon1 ,pterodactylus1 and       
,rhamphorhynchus that the resemblance is
 closest with the slender carnivorous   
,dinosaurs4 ,they never have the head of
 the thigh bone1 femur1 separated from  
its shaft by a constricted neck1 as in  
the ,pterodactyles from the ,chalk4 ,in 
many ways the thigh bone of ,dinosaurs  
tends towards being ,avian2 while that  
of ,pterodactyles inclines towards being
 ,mammalian1 but with a tendency to be  
,bird-like in the older types1 and to be

 ,mammal-like in the most recent        
representatives of the group in the     
,chalk4  ,the bones of the leg in       
,ornithosaurs1 known as tibia and       
fibula1 are remarkable for the          
circumstance first that they resemble   
,birds in the fibula being slender and  
only developed in its upper part towards
 the femur1 and secondly that in a genus
 like ,dimorphodon this drum-stick bone 
has the two upper bones of the ankle    
blended with the tibia1 so as to form a 
rounded pulley joint which is           
indistinguishable from that of a ,bird  
7see p4 #ajb74 ,there is a large number 
of ,dinosaurs in which this remarkable  
distinctive character of ,birds is also 
found4 ,only1 ,dinosaurs like           
,iguanodon1 for instance1 have the      
slender fibula as long as the tibia1 and
 contributing to unite with the separate
 ankle bones of the similarly rounded   
pulley at the lower end4 ,there are no  
,birds in which the tarsal bones remain 
separated and distinct throughout life4 

,but in ,pterodactylus from ,solenhofen1
 as in a number of ,dinosaurs1          
especially the carnivorous genera1 the  
bones of the tarsus remain distinct     
throughout life1 and never acquired such
 forms as would have enabled the ankle  
bone1 termed astragalus1 to embrace the 
extremity of the tibia1 as it does in   
,iguanodon4 ,thus the resemblance of the
 ,ornithosaur drum-stick is almost as   
close to ,dinosaurs as to ,birds4       
,there is great similarity between      
,dinosaurs and ,pterodactyles seen in   
the region of the instep1 known as the  
metatarsus4 ,these bones are usually    
four in number1 parallel to each other1 
and similar in form4 ,they are commonly 
longer than in ,dinosaurs2 but among    
some of the carnivorous ,dinosaurs their
 length approximates to that seen in    
,pterodactyles4 ,in neither group are   
the bones blended together by bony      
union1 while they are always united in  
,birds1 as in ,oxen and similar         
even-hoofed mammals4 ,dinosaurs agree   

with ,pterodactyles in maintaining the  
metatarsal bones separate1 but they     
differ from them and agree with ,birds  
frequently1 in having the number of     
metatarsal bones reduced to three1 as in
 ,iguanodon1 though ,dinosaurs often    
have as many as five digits developed4  
,the toe bones1 the phalanges of these  
digits of the hind limb1 are usually    
longer in ,pterodactyles than in        
,dinosaurs1 but they resemble           
carnivorous ,dinosaurs in the forms of  
their sharp terminal bones for the      
claws1 which are similarly compressed   
from side to side4  ,so diverse are the 
functions of the fore limb in ,dinosaurs
 and ,pterodactyles1 and so remarkably  
does the length of the metacarpal region
 of the back of the hand vary in the    
long-tailed and short-tailed            
,ornithosaurs1 that there is necessarily
 a less close correspondence in that    
region of the skeleton between these two
 groups of animals2 for the             
,pterodactyle fore limb is modified in  

relation to a function which can only be
 paralleled among ,birds and ,bats2 and 
yet neither of those groups of animals  
approximates closely in this region of  
the skeleton to the ,flying ,reptile4   
,under all the modifications of         
structure which may be attributed to    
differences of function1 some           
resemblance to ,dinosaurs may be        
detected1 which is best evident in the  
upper arm bone1 humerus2 is slight in   
the fore-arm bones1 ulna and radius2 and
 becomes lost towards the extremity of  
the limb4  ,if the tendency of the thigh
 bone to resemble a ,mammalian type of  
femur 7p4 #ajj7 is a fundamental1       
deep-seated character of the skeleton1  
it might be anticipated that a trace of 
,mammalian character would also be found
 in the humerus4 ,for what the character
 is worth1 the head of the humerus does 
show a closer approximation to a        
,monotreme ,mammal than is seen in      
,birds1 and is to some extent paralleled
 in those ,south ,african reptiles which

 approximate to ,mammals most closely4  
,not the least remarkable of the many   
astonishing resemblances of these light 
aerial creatures to the more heavy      
bodied ,dinosaurs is the circumstance   
that the humerus in both groups makes a 
not dissimilar approach to that of      
certain ,mammals4  ,these illustrations 
may be accepted as demonstrating a      
relationship between the ,ornithosaurs  
and ,dinosaurs now compared1 which can  
only be explained as results of         
influence of a common parentage upon the
 forms of the bones4 ,but more          
interesting than resemblances of that   
kind is the similarity that may be      
traced in the way in which air is       
introduced into cavities in the bones in
 both groups4 ,in some of the           
imperfectly known ,dinosaurs1 like      
,aristosuchus1 ,coelurus1 and           
,thecospondylus1 the bone texture is as 
thin as in ,pterodactyles1 and the      
vertebrae are excavated by pneumatic    
cavities1 which are amazing in size when

 compared with the corresponding        
structures in birds1 for the vertebra is
 often hollowed out so that nothing     
remains but a thin external film like   
paper for its thickness4 ,in the        
,dinosaurian genus ,coelurus this       
condition is as well marked in the tail 
and back as it is in the neck4 ,the     
essential difference from ,birds appears
 to be that in the larger carnivorous   
,dinosaurs the pneumatic condition of   
the bones is confined to the vertebral  
column2 while ,birds and ,pterodactyles 
have the pneumatic condition more       
conspicuously developed in the limb     
bones4 ,the pneumatic skeleton1 however1
 appears to be absent from the          
herbivorous types like ,iguanodon and   
all ,dinosaurs which have the ,bird-like
 form of pelvis1 and are most ,bird-like
 in the forms of bones of the hind limb4
 ,it is possible that some of the       
carnivorous ,dinosaurs also possessed   
limb bones with pneumatic cavities4     
,many of those bones are hollow with    

very thin walls4 ,if their cavities were
 connected with the lungs the foramina  
are inconspicuous and unlike the immense
 holes seen in the sides of the         
vertebrae4  ,according to the late      
,professor ,marsh1 the limbs of         
,coelurus and its allies1 which at      
present are imperfectly known1 are in   
some cases pneumatic4 ,therefore there  
is a closer fundamental resemblance     
between some carnivorous ,dinosaurs and 
,pterodactyles than might have been     
anticipated4 ,but the skull of ,coelurus
 is unknown1 and the fragments of the   
skeleton hitherto published are         
insufficient to do more than show that  
the two types were near in kindred1     
though distinct in habit4 ,each has     
elaborated a skeleton which owes much to
 the common stock which transmitted the 
vital organs1 and the tendency of the   
bones to take special forms2 but which  
also owes more than can be accurately   
measured to the action of muscles in    
shaping the bones and the influence of  

the mechanical conditions of daily life 
upon the growth of the bones in both of 
these orders of animals4 ,enough is     
known to prove that all ,dinosaurs      
cannot be regarded as ,ornithosaurs     
which have not acquired the power of    
flight2 though the evidence would lead  
us to believe that the primitive        
,ornithosaur was a four-footed animal1  
before the wing finger became developed 
in the fore limb as a means of extending
 a patagial membrane1 like the membrane 
which in the hind limb of ,dimorphodon  
has bent the outermost digit of the foot
 upward and outward to support the      
corresponding organ of flight extending 
down the hind legs4  ,it may thus be    
seen that the characters of             
,ornithosaurs which have already been   
spoken of as ,reptilian1 as             
distinguished from the resemblances to  
,birds1 may now with more accuracy be   
regarded as ,dinosaurian4 ,the          
,dinosaurs1 like ,pterodactyles1 must be
 regarded as intermediate in some       

respects between ,reptiles and ,birds4  
,the resemblances enumerated would alone
 constitute a partial transition from   
the ,reptile to the ,bird1 although no  
,dinosaurs have organs of flight2 many  
are heavily armoured with plates of     
bone1 and few1 if any1 approximate in   
the technical parts of the skeleton to  
the ,bird class1 except in the hind     
limbs4 ,yet ,dinosaurs have sometimes   
been regarded as standing to ,birds in  
the relation of ancestors1 or as        
parallel to an ancestral stock4  ,before
 an attempt can be made to estimate the 
mutual relation of the ,flying ,reptiles
 to ,dinosaurs on the one hand1 and to  
,birds on the other1 it may be well to  
remember that the resemblance of such a 
,dinosaur as ,iguanodon to a ,bird in   
its pelvis and hind limb is not more    
remarkable than that of ,pterodactyles  
to ,birds in the shoulder-girdle and    
bones of the fore limb4 ,the keeled     
sternum1 the long1 slender coracoid     
bones and scapulae1 are absolutely      

,bird-like in most ,ornithosaurs2 and   
that region of the skeleton only differs
 from ,birds in the absence of a        
furculum which represents the clavicles1
 and is commonly named the              
merry-thought4 ,the elongated bones of  
the fore-arm and the hand1 terminating  
in three sharp claws1 are characters in 
which the fossil bird ,archaeopteryx    
resembles the ,pterodactyle             
,rhamphorhynchus1 a resemblance which   
extends to a similar elongation of the  
tail4 ,it is remarkable that the        
resemblance should be so close1 since   
,archaeopteryx affords the only bird's  
skeleton known to be contemporary which 
can be compared with the ,solenhofen    
,flying ,reptiles4 ,the resemblance may 
possibly be closer than has been        
imagined4 ,the back of the head of      
,archaeopteryx is imperfectly preserved 
in the region of the quadrate bone1     
malar arch1 and temporal vacuity4 ,and  
till these are better known it cannot be
 affirmed that the back of the head is  

more ,reptilian in ,pterodactyles than  
in the oldest ,birds4 ,the side of the  
head in ,archaeopteryx is distinguished 
by the nostril being far forward1 the   
vacuity in front of the orbit being as  
large as in the ,pterodactyle           
,scaphognathus from ,solenhofen and     
other long-tailed ,pterodactyles4       
,,chapter ,,xviii  ,,how ,,pterodactyles
 ,,may ,,have ,,originated              
,ornithosauria have many characters     
inseparably blended together which are  
otherwise distinctive of ,reptiles1     
,birds1 and ,mammals1 and associated    
with peculiar structures which are      
absent from all other animals4 ,they are
 not quite alone in this incongruous    
combination of different types of       
animals in the same skeleton4           
,dinosaurs1 which were contemporary with
 ,ornithosaurs1 approximate to them in  
blending characters of ,birds with the  
structure of a ,reptile and something of
 a ,mammal in one animal4 ,if an        
,ornithosaur is ,reptilian in its       

backbone1 in the articular ends of each 
vertebra having the cup in front and    
ball behind in the manner of            
,crocodiles1 ,serpents1 and many        
,lizards1 a ,dinosaur like ,iguanodon1  
which had the reversed condition of ball
 in front and cup behind in its early   
vertebrae1 may be more ,mammalian than  
,avian in a corresponding resemblance of
 the bones to the neck in hoofed        
,mammals4 ,but while ,pterodactyles are 
sometimes ,mammalian in having the head 
of the thigh bone moulded as in         
carnivorous ,mammals and ,man1 the      
corresponding bone in a ,dinosaur is    
more like that of a ,bird4 ,and while   
the ,pterodactyle shoulder-girdle is    
often absolutely ,bird-like1 that region
 in ,dinosaurs can only be paralleled   
among ,reptiles4  ,such combinations of 
diverse characters are not limited to   
animals which are extinct4 ,there were  
not wanting scientific men who regarded 
the ,platypus of ,australia1 when first 
sent to ,europe1 as an ingenious example

 of ,eastern skill1 in which an animal  
had been compounded artificially by     
blending the beak of a ,bird with the   
body of a ,mammal4 ,fuller knowledge of 
that remarkable animal has continuously 
intensified wonder at its combination of
 ,mammal1 ,bird1 and ,reptile in a      
single animal4 ,it has broken down the  
theoretical divisions between the higher
 ,vertebrata1 demonstrating that a      
,mammal may lay eggs like a ,reptile or 
,bird1 that the skull may include the   
reptilian characters of the malar arch  
and pre-frontal and post-frontal bones1 
otherwise unknown in ,mammals and       
,birds4 ,the groups of ,mammals1 ,birds1
 and ,reptiles now surviving on the     
earth prove to be less sharply defined  
from each other when the living and     
extinct types are considered together4  
,but in ,pterodactyles1 ,mammal ,bird   
and ,reptile lose their identity1 as    
three colours would do when unequally   
mixed together4  ,this mingling of      
characteristics of different animals is 

not to be attributed to interbreeding1  
but is the converse of the combination  
of characters found in hybrid animals4  
,it is no exaggeration to say that there
 is a sense in which ,mammal1 ,bird1    
,reptile1 and the distinctive structures
 of the ,ornithosaur1 have              
simultaneously developed from one egg1  
in the body of one animal4  ,the        
differences between those vertebrate    
types of animals consist chiefly in the 
way in which their organisation is      
modified1 by one strain of characters   
being eliminated so that another becomes
 predominant1 while a distinctive set of
 structures is elaborated in each class 
of animals4 ,the earlier geological     
history of the higher ,vertebrata is    
very imperfectly known1 but the evidence
 tends to the inference that the older  
representatives of the several classes  
approximate to each other more closely  
than do their surviving representatives1
 so that in still earlier ages of time  
the distinction between them had not    

become recognisable4 ,the relation of   
the great groups of animals to each     
other1 among ,vertebrata1 is essentially
 a parallel relation1 like the colours  
of the solar spectrum1 or the parallel  
digits of the hand4 ,it was natural1    
when only the surviving life on the     
earth was known1 to imagine that animals
 were connected in a continuous chain by
 successive descent1 but ,mammals have  
given no evidence of approximation to   
,birds2 and ,birds discover no evidence 
that their ancestors were ,reptiles1 in 
the sense in which that word is used to 
define animals which now exist on the   
earth4 ,when the variation which animals
 attain in their maturity and exhibit in
 development from the egg was first     
realised1 it was imagined that ,nature1 
by slow summing up and accumulation of  
differences which were observed1 would  
so modify one animal type that it would 
pass into another4 ,there is little     
evidence to support belief that the     
changes between the types of life have  

been wrought in that way4 ,the history  
of fossil animals has not shown         
transitions of this kind from the lower 
to higher ,vertebrata1 but only         
intermediate1 parallel groups of        
animals1 analogous to those which       
survive1 and distinct from them in the  
same way as surviving groups are        
distinct from each other4 ,the          
circumstance that ,mammals1 ,birds1 and 
,reptiles are all known low down in the 
,secondary epoch of geological time1 is 
favourable to the idea of their history 
being parallel rather than successive4  
,such a conception is supported by the  
theory of elimination of characters from
 groups of animals as the basis of their
 differentiation4 ,this loss appears    
always to be accompanied by a           
corresponding gain of characters1 which 
is more remarkable in the soft1 vital   
organs than in the skeleton4 ,the gain  
in higher ,vertebrates in the bones is  
chiefly in the perfection of joints at  
their extremities2 but the gain in      

brain1 lungs1 heart1 and other soft     
parts is an elaboration of those        
structures and an increase in amount of 
tissue4  ,the resemblances of           
,ornithosaurs to ,mammals are the least 
conspicuous of their characters4 ,those 
seen in the upper arm bone and thigh    
bone are manifestly not derived from    
,mammals4 ,they cannot be explained as  
adaptations of the bones to conditions  
of existence1 because there is no       
community of habit to be inferred       
between ,pterodactyles and ,mammals1 in 
which the bones are in any way          
comparable4  ,other fossil animals show 
that a fundamentally ,reptilian         
structure is capable of developing in   
the ,mammalian direction in the skull1  
backbone1 shoulder-girdle1 hip-girdle1  
and limbs1 so as to be uniformly        
,mammalian in its tendencies4 ,this is  
proved by tracing the ,north ,american  
,texas fossils named ,labyrinthodonts1  
through the ,south ,african             
,theriodonts1 towards the ,monotremata  

and other ,mammalia4 ,just as those     
animals have obliterated all traces of  
the ,bird from their skeletons1 ,birds  
have obliterated the distinctive        
characters of ,mammals4 ,the            
,ornithosaur has partially obliterated  
both4 ,with a skull and backbone marked 
by typical characters of the ,reptile1  
it combines the shoulder-girdle and     
hip-girdle of a ,bird1 with characters  
in the limbs which suggest both those   
types in combination with ,mammals4     
,the bones have been compared in the    
skeleton of each order of existing      
,reptiles1 and found to show side by    
side with their peculiar characters not 
only resemblances to the other          
,reptilia1 but an appreciable number of 
,mammalian and ,avian characters in     
their skeletons4 ,the term crocodile1   
for example1 indicates an animal in     
which the skeleton is dominated by one  
set of peculiar characters4 ,crocodiles 
retain enough of the characteristics of 
several other orders of reptiles to show

 that an animal sprung from the old     
,crocodile stock might diverge widely   
from existing ,crocodiles by            
intensifying what might be termed its   
dormant characters in the ,crocodile    
skeleton4 ,comparing animals together   
bone by bone it is possible to value the
 modifications of form which they put   
on1 and the resemblances between them1  
so as to separate the inherited wealth  
of an animal's affinities with ancestors
 or collateral groups1 from the peculiar
 characters which have been acquired as 
an increase based upon its typical bony 
possessions or osteological capital4    
,there is no part of the ,pterodactyle  
skeleton which is more distinctly       
modified than the head of the upper arm 
bone1 which fits into the socket between
 the coracoid bone and the              
shoulder-blade4 ,the head of the        
humerus1 as the articular part is named1
 is somewhat crescent-shaped1 convex on 
its inner border1 and a little concave  
on its outer border1 and therefore      

unlike the ball-shaped head of the upper
 arm bone in ,man and the higher        
,mammals4 ,it is much more nearly       
paralleled in the little group of       
,monotremata allied to the living       
,ornithorhynchus4 ,in that sense the    
head of the humerus in a ,pterodactyle  
has some affinity with the lowest       
,mammalia1 which approach nearest to    
,reptiles4 ,the character might pass    
unregarded if it were not found in more 
striking development in fossil ,reptiles
 from ,cape ,colony1 which from having  
teeth like ,mammals are named           
,theriodontia4 ,in several of those     
,south ,african reptiles the upper arm  
bone approaches closer to the humerus in
 ,ornithosaurs than to ,ornithorhynchus4
 ,such coincidences of structure are    
sometimes dismissed from consideration  
and placed beyond investigation by being
 termed adaptive modifications2 but     
there can be no hope of finding         
community of habit between the burrowing
 ,monotreme1 the short-limbed           

,theriodont1 and the flying             
,pterodactyle which might have caused   
this articular part of the upper arm    
bone to acquire a form so similar in    
animals constructed so differently4 ,if 
the resemblance in the humerus to       
,monotremes in this respect is not to be
 attributed to burrowing1 neither can   
the crescent form of its upper          
articulation be attributed to flight2   
for in ,birds the head of the bone is   
compressed1 but always convex1 and ,bats
 fly without any approach to the        
,pterodactyle form in the head of the   
humerus4 ,this apparently trivial       
character may from such comparisons be  
inferred to be something which the way  
of life of the animal does not          
sufficiently account for4 ,these        
deepest-seated parts of the limbs are   
slow to adapt themselves to changing    
circumstances of existence1 and retain  
their characters with moderate variation
 of the bones in each of the orders or  
classes of animals4 ,it therefore is    

safer to regard ,mammalian characters1  
as well as the resemblances which       
,pterodactyles show to other kinds of   
animals1 as due to inheritance from a   
time when there was a common stock from 
which none of these animals which have  
been considered had been distinctly     
elaborated4  ,a few characters of       
,ornithosaurs are regarded as having    
been acquired1 because they are not     
found in any other animals1 or have been
 developed only in a portion of the     
group4 ,the most obvious of these is the
 elongated wing finger2 but in some     
genera1 like ,dimorphodon1 there is also
 a less elongation of the fifth digit of
 the foot1 and perhaps in all genera    
there is a backward development of the  
first digit of the hand1 which is       
without a claw1 and therefore unlike the
 clawed digit of a ,bat4 ,an acquired   
character of another kind1 which is     
limited to the ,cretaceous genera1 is   
seen in the shoulder-blade being        
directed transversely outward1 so that  

its truncated end articulates by a true 
joint with the early vertebrae of the   
back1 and defended the cavity inclosed  
by the ribs by a strong bony external   
arch4 ,and finally1 as the animals later
 in time acquire short tails1 and       
relatively longer limbs1 the bones of   
the back of the hand1 termed            
metacarpals1 acquire greater and        
distinctive length1 which is not seen in
 the long-tailed types like             
,rhamphorhynchus4  ,these and such-like 
acquired characters distinguish the     
class of animals from all groups with   
which it may be compared1 and mark the  
possible limits of variation of the     
skeleton within the boundary of the     
order4 ,but no further variation of     
these parts of the skeleton could make a
 transition to another order of animals1
 or explain how the ,pterodactyles came 
into existence1 because the characters  
which separate orders and classes of    
animals from each other differ in kind  
from those which separate smaller       

groups1 named genera and species1 of    
which the order is made up4 ,the        
accumulation of the characters of genera
 will not sum up into the characters of 
an order or class4  ,in making the      
division of ,vertebrate animals into    
classes the skeleton is often almost    
ignored4 ,its value is entirely         
empirical and based upon the observed   
association of the various forms of     
bones with the more important characters
 of the brain and other vital organs4   
,what is understood as a ,mammalian or  
,avian character in the skeleton is the 
form of bone which is found in          
association with the soft vital organs  
which constitute an animal a ,mammal or 
a ,bird4  ,the characters which         
theoretically define a ,mammal appear to
 be the enormous overgrowth of the      
cerebral hemispheres of the brain by    
which the cerebrum comes into contact   
with the cerebellum1 as among ,birds4   
,this character distinguishes both      
groups of animals from all ,reptiles1   

recent and fossil4 ,but in examining the
 mould of the interior of the brain case
 it is rare to have the bones fitting so
 closely to the brain as to prove that  
the lateral expansion below the cerebrum
 and cerebellum is formed by the optic  
lobes of the brain4 ,otherwise the brain
 of a ,pterodactyle might be as like to 
the brain of ,ornithorhynchus as it is  
like that of a ,bird 7,fig4 #ai74 ,but  
it is precisely in this condition of    
arrangement of the parts of the brain   
that the specimens appear to be most    
clear4 ,the lateral mass of brain in    
specimens of ,ornithosaurs from the     
,lower ,secondary rocks appears to be   
transversely divided into back and front
 parts1 which may be thought to         
correspond to the structures in a       
,mammal brain named corpora             
quadrigemina1 but to be placed as the   
optic lobes are placed in ,birds1 and to
 have relatively greater dimensions than
 in ,mammals4 ,no evidence has been     
observed of this transverse division of 

the optic lobes of the brain in         
,pterodactyles from the ,chalk and      
,cretaceous rocks1 and so far as the    
evidence goes this part of the brain was
 shaped as in birds1 but rather smaller4
  ,the brain is the only soft organ in  
which a ,mammalian character could be   
evidenced4 ,the uniformity in character 
of the brain throughout the group in    
,mammals is remarkable1 in reference to 
the circumstance that the reproduction  
varies in type2 the lowest1 or          
,monotreme division1 being oviparous4   
,if there is no necessary connexion     
between the ,mammalian brain and the    
prevalent condition under which the     
young are produced alive1 it may be     
affirmed also that there is no necessary
 connexion between the form of the brain
 and the form of the bones1 since the   
brain cavity in ,theriodont reptiles    
shows no resemblance to that of a       
,mammal1 while the bones are in so many 
respects only paralleled among          
,monotremata and ,mammalia4 ,the variety

 of forms which the existing ,mammalian 
orders of animals assume1 shows the     
astonishing range of structure of the   
skeleton which may coexist with the     
,mammalian brain4 ,and therefore we are 
led to the conclusion that any other    
fundamental modification of brain--such 
as distinguishes the class of           
,birds--might also be associated with   
forms and structures of the skeleton    
which would vary in similar ways4 ,in   
other words1 if for convenience we      
define a ,mammal by its form of brain1  
structure of the heart and lungs1 and   
provision for nutrition of the young1   
without regard to the covering of the   
skin1 which varies between the scales of
 a pangolin and the practically naked   
skin of the whale--a bird might be also 
defined by its peculiar conditions of   
brain and lungs1 without reference to   
the feathered condition of the skin1    
though the feathered condition extends  
backward in time to the ,upper          
,secondary rocks1 as seen in the        

,archaeopteryx4  ,the ,avian characters 
of ,pterodactyles are the predominant   
parts of their organisation1 for the    
conditions of the brain and lungs shown 
by the moulds of the brain case and the 
thin hollow bones with conspicuous      
pneumatic foramina1 give evidence of a  
community of vital structures with      
,birds1 which is supported by characters
 of the skeleton4 ,if any               
classificational value can be associated
 with the distribution of the pneumatic 
foramina as tending to establish        
membership of the same class for animals
 fashioned on the same plan of soft     
organs1 the evidence is not weakened    
when a community of structures is found 
to extend among the bones to such       
distinctive parts of the skeleton as the
 sternum1 shoulder-girdle1 bones of the 
fore-arm and fore-leg2 for in all these 
regions the ,pterodactyle bones are     
practically indistinguishable from those
 of ,birds4 ,this is the more remarkable
 because other parts of the skeleton1   

such as the humerus and pelvis1 show a  
partial resemblance to ,birds1 while the
 parts which are least ,avian1 like the 
neck bones1 have no tendency to vary the
 number of the vertebrae1 in the way    
which is common among ,birds1 following 
more closely the formula of the seven   
cervical vertebrae of ,mammals4  ,it    
would therefore appear from the vital   
community of structures with ,birds1    
that ,pterodactyles and ,birds are two  
parallel groups1 which may be regarded  
as ancient divergent forks of the same  
branch of animal life1 which became     
distinguished from each other by        
acquiring the different condition of the
 skin1 and the structures which were    
developed in consequence of the bony    
skeleton ministering to flight in       
different ways2 and with different habit
 of terrestrial progression1 this       
extinct group of animals acquired some  
modifications of the skeleton which     
,birds have not shown4 ,there is nothing
 to suggest that ,pterodactyles are a   

branch from ,birds1 but their relation  
to ,birds is much closer1 so far as the 
skeleton goes1 than is their relation   
with the flightless ,dinosaurs1 with    
which ,birds and ,pterodactyles have    
many characters in common4  ,on the     
theory of elimination of character which
 ,i have used to account for the        
disappearance of some ,mammalian        
characters from the ,pterodactyle1 that 
loss is seen chiefly in the removal of  
the parts which have left a ,reptilian  
articulation of the lower jaw with the  
skull1 and the articulation of the      
vertebrae throughout the vertebral      
column by a modified cup-and-ball form  
of joint4 ,the furculum of the ,bird is 
always absent from the ,pterodactyle4   
,no specimen has shown recognisable     
clavicles or collar-bones4 ,judged by   
the standard of existing life1          
,pterodactyles belong to the same group 
as ,birds1 on the evidence of brain and 
lungs1 but they belong to a different   
group on account of the dissimilar      

modifications of the skeleton and       
apparent absence of feathers from the   
skin4  ,the most impressive facts in the
 ,pterodactyle skeleton1 in view of     
these affinities1 are the structures    
which it has in common with ,reptiles4  
,some structures are fundamental1 like  
the cup-and-ball articulation of the    
vertebrae1 which is never found in birds
 or mammals4 ,although not quite        
identical with the condition in any     
,reptile1 this structure is             
approximately ,lizard-like or           
,crocodile-like in the cup-and-ball     
character4 ,it shows that the           
deepest-seated part of the skeleton is  
,reptile-like1 though it may not be more
 ,reptilian than is the vertebral column
 of a ,mammal1 if comparison is made    
between ,mammals and extinct groups of  
animals known as ,reptiles1 such as     
,dinosaurs and ,theriodontia4  ,the     
orders of animals which have been       
included under the name ,reptilia       
comprise such different structural      

conditions of the parts of the skeleton 
which may be termed reptilian in        
,ornithosaurs1 that there is good reason
 for regarding the cup-and-ball         
articulation as quite a distinctive     
,reptilian specialisation1 in the same  
sense that the saddle-shaped            
articulation between the bodies of      
adjacent vertebrae in a bird is an      
,avian specialisation4 ,from the        
theoretical point of view the           
,ornithosaur acquired its ,reptilian    
characters simultaneously with its      
,avian and ,mammalian characters4       
,there is nothing in the structure of   
the skeleton of the ,dinosauria1 to     
which ,ornithosaurs approximate in      
several parts of the body1 which would  
help to explain the cup-and-ball        
articulation of the backbone1 if the    
,flying ,reptile were supposed to be an 
offshoot from the carnivorous           
,dinosaurs4  ,the elimination of        
,reptile characters from so much of the 
skeleton1 and the substitution for them 

of the characters of ,birds and         
,mammals1 would be of exceptional       
interest if there had been any ground   
for regarding the flying animal as more 
nearly related to a ,reptile than to a  
,bird4 ,but if the evidence from the    
form of the brain and nature of the     
pneumatic organs seen in the limb bones 
accounts for the ,avian features of the 
skeleton1 the ,reptilian condition of   
the vertebral column helps to show a    
capacity for variation1 and that the    
fixity of type and structure1 which the 
skeleton of the modern ,bird has        
attained1 is not necessarily limited to 
or associated with the vital organs of  
,birds4  ,the variation of the          
cup-and-ball articulation in the neck of
 a ,chelonian1 which makes the third    
vertebra cupped behind1 the fourth      
bi-convex1 the fifth cupped in front1   
and the sixth flattened behind1 shows   
that too much importance may be attached
 to the mode of union of these bones in 
,serpents1 ,crocodiles1 and those       

,lizards which have the cup in front2   
for while in ,lizards the anterior cup1 
oblique and depressed1 is found in most 
of its groups1 the ,geckos show no trace
 of the cup-and-ball structure1 and in  
that respect resemble the ,hatteria of  
,new ,zealand4  ,if1 therefore1 the     
cup-and-ball articulation of vertebrae  
in ,ornithosauria has any significance  
as a mark of affinity to ,reptiles1 it  
could only be in approximation to those 
living ,reptiles which possess the same 
character1 and would have it on the     
hypothesis that both have preserved the 
structure by descent from an earlier    
type of animal4 ,this hypothesis is     
negatived by the fact that the          
cup-and-ball articulation is unknown in 
the older fossil ,reptiles4  ,although  
the articulation for the lower jaw with 
the skull in ,ornithosaurs is only to be
 paralleled among ,reptiles1 the        
structure is adapted to a brain case    
which is practically indistinguishable  
from that of a ,bird1 except for the    

postorbital arch4  ,the hypothesis of   
descent1 therefore1 becomes impossible1 
in any intelligible form1 in explanation
 of distinctive character of the        
skeleton4 ,the hypothesis of elimination
 may also seem to be insufficient1      
unless the potential capacity for new   
development be recognised as concurrent1
 and as capable of modifying each region
 of the skeleton1 or hard parts of the  
animal1 in the same way that the soft   
organs may be modified4 ,from which we  
infer that all structures1 which        
distinguish the several grades of       
organisation in modern classifications1 
soft parts and hard parts alike1 may    
come into existence together1 in so far 
as they are compatible with each other1 
in any class or ordinal division of     
animals4  ,although the young ,mammal   
passes through a stage of growth in     
which the brain may be said to be       
,reptilian1 there is no good ground for 
inferring that ,mammal or ,bird type of 
skeleton was developed later in time    

than that of ,reptiles4 ,the various    
types of ,fishes have the brains in     
general so similar to those of ,reptiles
 that it is more intelligible for all   
the vertebrate forms of brain to have   
differentiated at the same time1 under  
the law of elimination of characters1   
than that there should be any other bond
 of union between the classes of        
animals4  ,if we ask what started the   
,ornithosauria into existence1 and      
created the plan of construction of that
 animal type1 ,i think science is       
justified in boldly affirming that the  
initial cause can only be sought under  
the development of patagial membranes1  
such as have been seen in various       
animals ministering to flight4 ,such    
membranes1 in an animal which was       
potentially a ,bird in its vital organs1
 have owed development to the absence of
 quill feathers4 ,thus the wing membrane
 may be the cause for the chief         
differences of the skeleton by which    
,ornithosaurs are separated from ,birds1

 for the stretch of wing in one case is 
made by the skin attached to the bones1 
and in the other case by feathers on the
 skin so attached as to necessitate that
 the wing bones have different          
proportions from ,ornithosaurs4  ,it is 
a well-known observation that each great
 epoch of geological time has had its   
dominant forms of animal life1 which1 so
 far as the earth's history is known    
now1 came into existence1 lived their   
time1 and were seen no more4 ,in the    
same way the smaller groups of species  
and genera included in an ordinal group 
of animals or class have abounded1      
giving a tone to the life of each       
geological formation1 until the vitality
 of the animal is exhausted1 and the    
species becomes extinct or ceases to    
preponderate4 ,this process is seen to  
be still modifying the life on the      
earth1 when some kinds of animals and   
plants are introduced to new conditions4
 ,plants appear to wage successful war  
more easily than animals4 ,the          

introduction of the ,cactus in some     
parts of ,cape ,colony has locally      
modified both the fauna and flora1 just 
as the ,anacharis introduced into       
,england spread from ,cambridge over the
 whole country1 and became for many     
years the predominant form of plant life
 in the streams4 ,the ,rabbit in        
,australia is a historic pest4          
,something similar to this physical     
fertility and increase appears to take  
place under new circumstances in certain
 organs within the bodies of animals1 by
 the development of structures          
previously unknown4 ,a familiar example 
is seen in the internal anatomy of the  
,trout introduced into ,new ,zealand1   
where the number of pyloric appendages  
about the stomach has become rapidly    
augmented1 while the size and the form  
of the animal have changed4 ,the        
rapidity with which some of these       
changes have been brought about would   
appear to show that ,nature is capable  
of transforming animals more rapidly    

than might have been inferred from their
 uniform life under ordinary            
circumstances4 ,growth of the vital     
organs in this way may modify the       
distinctive form of any vital organ1    
brain or lungs1 and thus as a           
consequence of modification of the      
internal structures due to changes of   
food and habit1 bring a new group of    
animals into existence4 ,and just as the
 group of animals ceases to predominate 
after a time1 so there comes a limit to 
the continued internal development of   
vital structures as their energy fails1 
for each organ behaves to some extent   
like an independent organism4  ,under   
such explanations of the mutual         
relations of the parts of animals1 and  
groups of animals1 time ceases to be a  
factor of primary importance in their   
construction or elaboration4 ,the       
supposed necessity for practically      
unlimited time to produce changes in the
 vital organs which separate animals    
into great orders or classes is a       

nightmare1 born of hypothesis1 and may  
be profitably dismissed4 ,the geological
 evidence is too imperfect for dogmatism
 on speculative questions2 but the      
nature of the affinities of             
,ornithosaurs to other animals has been 
established on a basis of comparison    
which has no need of theory to justify  
the facts4 ,it is not improbable that   
the primary epoch of time1 even as known
 at present1 may be sufficiently long to
 contain the parent races from which    
,ornithosaurs and all their allies have 
arisen4  ,in thus stating the relation  
of ,ornithosaurs to other animals the   
,flying ,reptile has been traced home to
 kindred1 though not to its actual      
parents or birthplace4 ,there is no     
geological history of the rapid or      
gradual development of the wing finger1 
and although the wing membrane may be   
accepted as its cause of existence1 the 
wing finger is powerfully developed in  
the oldest known ,pterodactyles as in   
their latest representatives4           

,pterodactyles show singularly little   
variation in structure in their         
geological history4 ,we chronicle the   
loss of the tail and loss of teeth4     
,there is also the loss of the outermost
 wing digit from the hind foot as a     
supporter of the wing membrane4 ,but the
 other variations are in the length of  
the metacarpus1 or of the neck1 or head4
 ,one of the fundamental laws of life   
necessitates that when an animal type   
ceases to adapt its organisation and    
modify its structures to suit the       
altered circumstances forced upon it by 
revolutions of the earth's surface its  
life's history becomes broken4 ,it must 
bend or break4  ,the final disappearance
 of these animals from the earth's      
history in the ,chalk may yet be        
modified by future discoveries1 but the 
,flying ,reptiles have vanished1 in the 
same way as so many other groups of     
animals which were contemporary with    
them in the ,secondary period of time4  
,such extinctions have been attributed  

to catastrophes1 like the submergence of
 land1 so that the habitations of       
animals became an area gradually        
decreasing in size1 which at last       
disappeared4 ,it appears also to be a   
law of life1 illustrated by many extinct
 groups of animals1 that they endure for
 geological ages1 and having fought     
their battle in life's history1 grow old
 and unable to continue the fight1 and  
then disappear from the earth1 giving   
place to more vigorous types adapted to 
live under new conditions4  ,the extinct
 ,pterodactyles hold a relation to      
,birds in the scheme of life not unlike 
that which ,monotremata hold to other   
,mammals4 ,both are remarkable for the  
variety of their affinities and         
resemblances to ,reptiles4 ,the         
,ornithosauria have long passed away2   
the ,monotremes are nearing extinction4 
,both appear to be supplanted by        
parallel groups which were their        
contemporaries4 ,birds now fill the     
earth in a way that ,flying ,reptiles   

never surpassed2 but their flight is    
made in a different manner1 and the wing
 is extended to support the animal in   
the air1 chiefly by appendages to the   
skin4  ,if these fossils have taught    
that ,ornithosaurs have a community of  
soft vital organs with ,dinosaurs and   
,birds1 they have also gone some way    
towards proving that causes similar to  
those which determined the structural   
peculiarities of their bony framework1  
originated the special forms of         
respiratory organs and brain which      
lifted them out of association with     
existing ,reptiles4   ,these old flying 
animals sleep through geological ages1  
not without honour1 for the study of    
their story has illuminated the mode of 
origin of animals which survive them1   
and in cleaving the rocks to display    
their bones we have opened a new page of
 the book of life4     ,,appendix   ,the
 best public collections of             
,ornithosaurian remains in ,england are 
in the ,british ,museum 7,natural       

,history72 ,museum of ,practical        
,geology1 ,royal ,college of ,surgeons2 
the ,university ,museum1 ,oxford2       
,geological ,museum1 ,cambridge2 and the
 ,museum of the ,philosophical ,society 
at ,york4  ,detailed descriptions and   
original figures of the principal       
specimens mentioned or referred to may  
be found in the following writings3--   
,h4 v4 ,meyer1 ,reptilien aus dem       
,lithograph4 ,schiefer4 #ahei4 ,folio4  
v4 ,quenstedt1 ,pterodactylus suevicus4 
#ahee4 #dto4   ,goldfuss1 ,nova ,acta   
,leopold4 ,,xv4   v4 ,munster1 ,nova    
,acta ,leopold4 ,,xv4   ,a4 ,wagner1    
,abhandl4 ,bayerischen ,akad41 vi41     
viii4   ,cuvier1 ,annales du ,museum1   
xiii4 #ahji4          ,ossemens         
fossiles1 v4 #ahbd4   ,buckland1 ,geol4 
,trans41 ser4 #b1 iii4   ,r4 ,owen1     
,palaeontographical ,society4 #ahea1    
#ahei1 #ahfj1 #ahgj1 #ahgd4   ,k4 v4    
,zittel1 ,palaeontographica1 xxix4      
#ahhb4   ,t4 ,c4 ,winkler1 ,mus4 ,teyler
 ,archives4 #ahgd1 #ahhc4   ,oscar      

,fraas1 ,palaeontographica1 xxv4 #ahgh4 
,anton ,fritsch1 ,boehm4 ,gesell4       
,sitzber4 #ahha4   ,r4 ,lydekker1       
,catalogue of ,fossil ,reptilia in      
,british ,museum ,i4 #ahhh4   ,o4 ,c4   
,marsh1 ,amer4 ,jour4 ,science4 #ahhb1  
#ahhd4   ,s4 ,w4 ,williston1 ,kansas    
,university ,quarterly4 #ahic1 #ahif4   
,e4 ,t4 ,newton1 ,phil4 ,trans4 ,royal  
,soc4 #ahhh1 #ahid4   ,h4 ,g4 ,seeley1  
,ornithosauria4 #hvo4 #ahgj4            
,annals and ,mag4 ,natural ,hist4 #ahgj1
 #ahga1 #ahij1 #ahia4                   
,linn4 ,society4 #ahgd1 #ahge4          
,geol4 ,mag4 #ahha4   ,felix            
,pleininger1 ,palaeontographica4 #ahid1 
#aija4     ,,index   ,a    ,abdominal   
ribs1 #he1 #aed    ,accumulation of     
characters1 #bbj    ,acetabulum1 #ie    
,acquired characters1 #bai    ,adjacent 
land1 #acf    ,air cells1 #aj1 #dh      
,albatross1 #bc1 #cf1 #agf              
,alligator1 brain1 #ec2     pelvis1 #ih 
,american ,greensand1 #ahe    --        
ornithosaurs1 #hg1 #abf    ,amphibia1   

#d1 #aia    ,anabas1 #ag    ,anacharis1 
#bbg    ,anchisaurus1 #aii    ,angle of 
lower jaw1 #ge    ,ankle bones1 #ajc1   
#aie1 #bjg    ,anomodonts1 #aib         
,ant-eater of ,africa1 #adb2     ,india1
 #dj2     ,south ,america1 #dj1 #ahe    
,apteryx1 lungs1 #dh2     pelvis1 #ie   
,aquatic mammals1 #ada    ,aramis1      
scapular arch1 #aac    ,archaeopteryx1  
#eh1 #gf1 #ajd1 #acj1 #aig1 #baa        
,aristosuchus1 #abi1 #aij1 #bje1 #bji   
,armadillo1 #dj1 #ada    ,articulation  
of the jaw1 #ab1 #ge    ,ashwell1 #agg  
,atlantosaurus1 #bjb    ,atlas and axis1
 #hj1 #ha    ,aves1 #aij    ,avian      
characters1 #bbj1 #bbb   ,b             
,backbone1 #gh1 #hd    ,banz1 #adh      
,barbastelle1 #be    ,barrington1 #agg  
,barton1 #agg    ,bat1 #ch1 #aaj1 #aig2 
sternum of1 #ajg2     metacarpus1 #abh  
,bavaria1 #aef1 #ahe    ,beak1 horny1   
#gd1 #agh    ,bear1 skull of1 #ab2      
femur1 #ajj    ,bel and the ,dragon1 #ae
    ,belodon1 #bjb    ,bird1 #hj1 #aaj1 
#abj    -- resemblances1 #fc1 #fe1 #ga1 

#ie1 #ajb1 #ajh1 #aac1 #aai1 #abj1 #baa 
,bird-reptile1 #ahh    ,bird wing1 #abh1
 #acj    ,birds in flight1 #bb2     with
 teeth1 #gf    ,black-headed bunting1   
#dg    ,blainville1 ,d4 de1 #cj1 #aic   
,blood1 temperature of1 #ef    ,bohemia1
 #cd    ,bonaparte1 ,prince ,charles1   
#cj    ,bones of birds1 variation in1   
#da    -- of reptiles1 variation in1 #db
    -- about the brain1 #fi    -- in the
 back1 #hd    ,bone texture1 #ei1 #bji  
,bonn ,museum1 #cb1 #he1 #aef    ,brain 
and breathing organs1 #ee    ,brain     
cavity1 in birds and reptiles1 #eb2     
in mammals1 #bba1 #bbf2     in          
,solenhofen pterodactyles1 #ed1 #bbj    
,brazil1 #cd    ,breathing organs1 #h   
,bridgewater ,treatise1 #adc    ,british
 ,museum1 #acc1 #ahc    ,brixton1 ,isle 
of ,wight1 #ee1 #agd    ,buckland1      
,dean1 #adc1 #adh1 #bca    ,burrowing   
limb1 #ch   ,c    ,cactus1 #bbg         
,calamospondylus1 #bjc    ,cambridge    
,greensand1 #cc1 #hi1 #agf    --        
,museum1 #agg    ,camel1 #hc            

,campylognathus1 #fh1 #ga1 #ace2        
size of1 #adi    ,canary1 #dg           
,carnivorous dinosaurs1 #abi    ,carpus1
 #abb    ,caudal fin1 #ia1 #afa    --   
vertebrae1 #hi1 #ib1 #bjc    ,ceratodus1
 #d1 #e1 #i1 #ag    ,ceratosaurus1 #bjc1
 #bjd    ,cervical rib1 #ha    ,cetacea1
 #dj    ,cetiosaurus1 #aih1 #bjc        
,chalinolobus1 #be    ,chalk1           
pterodactyles in1 #acf2     of ,kansas1 
#ajc1 #acb    ,chameleon1 #ag1 #ea1 #gj2
     scapula1 #aab2     sternum1 #ajg   
,chameleonoidea1 #aia    ,cheek bones1  
#agh    ,chelonia1 #hf1 #aab1 #aic      
,chesterton1 #agg    ,chlamydosaurus1   
#ba    ,chrysochloris capensis1 #aba    
,classification1 #aib2     on pelvis    
characters1 #aie2     of dinosaurs1 #aih
    ,clavicles1 #aaa1 #aab    ,claw1    
#aje1 #aaf1 #ahc1 #bjh    ,coelurus1    
#bjc1 #bji    ,coldham ,common1 #agg    
,collar bone1 #aaa    ,collini1 #bg     
,comparison with dinosaurs1 #aih2     of
 pelvis1 #bjd1 #bjf2     of skulls1     
#aib1 #aii1 #bja    ,cope1 ,professor1  

#ca1 #cd    ,coracoid1 #aji1 #aab1 #aac 
,cordylomorpha1 #aia    ,cormorant1 #gj1
 #agd2     sternum1 #ajh    ,corpora    
quadrigemina1 #bba    ,crisp1 ,dr41 on  
pneumatic skeleton1 #dg    ,crocodile1  
characters of1 #bag2     heart1 #ef2    
lung1 #i2     shoulder-girdle1 #aaa2    
skull1 #df2     vertebrae1 #gi          
,crocodilia1 #aij    ,curlew1 #fh       
,cuvier1 #a1 #bg1 #bh1 #ed1 #gf1 #gg1   
#acj1 #bca    ,cycnorhamphus1 #gj1 #id1 
#aga1 #agc1 #bjd    ,cycnorhamphus      
,fraasii1 #hj1 #if1 #afi    -- suevicus1
 #afi1 #agj    ,cypselus1 #db   ,d      
,dacelo gigantea1 #fc    ,darwin1 #c    
,davy1 ,dr4 ,john1 #adb                 
,deuterosaurus1 #ig    ,dicynodon1 #bjj 
,dicynodon lacerticeps1 #ga    ,digits1 
of ostrich1 #bc2     of pterodactyle1   
#abh    ,digits with claws1 #acj2       
foot bones in1 #aje    ,dimorphodon1    
#fc1 #fd1 #ff1 #fg1 #gc1 #gd1 #hc1 #ij1 
#ajb1 #aac1 #adc1 #aib1 #aid1           
#aii1 #bja1 #bjf    ,dinosauria1 #f1    
#gg1 #hd1 #hg1 #ie1 #abi1 #add1 #aih1   

#bji    ,dinosaurs from ,lias1 #ace1    
#aib2     from ,elgin1 #bja1 #bjg2      
,stuttgart1 #bjb2     ,trias dinosaurs1 
#aii1 #bjj    ,diopecephalus1 #afh      
,diving birds1 #bc1 #hc1 #ajb           
,dolichosauria1 #aia    ,dolphin1 #ajg  
,doratorhynchus1 #agc    ,dorygnathus1  
#gd1 #adh    ,dragons1 #c1 #ae1 #ag     
,drumstick bone1 #ajc1 #aie    ,duck1   
#bb1 #hc   ,e    ,echidna1 #ge1 #gf1    
#ie1 #ajj    ,edentata1 #ahe            
,edentulous beak1 #aec    ,eichstaedt1  
#cb    ,elephant1 head of1 #df          
,enumeration of characters1 #bbc1 #bbe  
,ephesus1 winged figure1 #af            
,epiphysis to first phalange1 #abc      
,exocoetus1 #ah    ,extinctions1 #abi   
,eye hole1 #add2     sclerotic bones in1
 #fe   ,f    ,farren1 ,william1 #cd     
,femur1 #ajj    ,fibula1 #ajb1 #ahc1    
#bjf    ,fifth outer digit1 #acb2     in
 foot1 #ade    ,figure from temple at   
,ephesus1 #af    ,first phalange1 #aea  
,fish-eating crocodile1 #acg    ,flight1
 organs of1 #ag2     in bats1 #be       

,flying limb1 #ch    ,flying fishes1    
#ah1 #eg2     foxes1 #bf2     frogs1    
#ai1 #aig2     gecko1 #ba1 #bd2         
lizards1 #bj2     reptiles1 #cg1 #df2   
squirrel1 #bd    ,foot1 #ajd2     digits
 in1 #aje1 #adf    ,fore leg1 #ajb1 #bjf
    -- limb1 #ch1 #ajg1 #aaf1 #abj      
,four claws1 #adg    ,fox1 ,rev4 ,w41   
#ee1 #agd    ,fraas1 ,professor ,oscar1 
#agb1 #bca    ,frigate bird1 vertebrae  
of1 #hf1 #agd    ,frog1 lungs of1 #h    
,furculum1 #aad   ,g    ,gaudry1        
,professor ,a41 #ca    ,gavial1 #acf    
,gecko1 #ba1 #bc    ,genera1 comparison 
of1 #aib    ,geological distribution1   
#ahf    ,gills1 #d    ,giraffe1 #ch1 #ci
    ,glossy starling1 #dg    ,golden    
eagle1 #abj    -- mole1 #aba            
,goldfuss1 #cj1 #bca    ,granchester1   
#agg    ,great ant-eater1 #dj1 #ahe     
,guillemot1 #ajb    ,gull1 #bb   ,h     
,haarlem1 ,teyler ,museum at1 #cb       
,habits1 probable1 #acd1 #agf1 #aih     
,hairless skins1 #ada    ,hand in       
mammals1 #ch    ,harston1 #agg          

,haslingfield1 #agg    ,hastings1 #agd  
,hatteria lung1 #i1 #bg2     brain1 #ec2
     skull1 #gj1 #gg2     ribs1 #hf2    
a reptile type1 #ac    ,head1 characters
 of1 #gf    ,heidelberg ,museum1 #cb1   
#ed1 #aei    ,herpetomorpha1 #aia       
,heron1 #fe1 #agd    ,hesperornis1 #gf  
,hind foot1 #ajd1 #ace    -- limb1 #ic1 
#ii1 #aei1 #bjf    ,hip-girdle in whale 
tribe1 #ci1 #aei    ,homoeosauria1 #aia 
,horningsea1 #agg    ,horse1 metacarpus 
of1 #abg2     vertebrae of1 #gi         
,humerus1 #df1 #aag1 #bag    ,huxley1   
,professor1 #ca1 #hi1 #aed1 #ahh        
,hyo-mandibular arch1 #ac    ,hypothesis
 of descent1 #bbf    ,hyrax1 #aja   ,i  
,ichthyornis1 #gf    ,ichthyosaurus1 #f1
 #aia    ,iguanodon1 #bji2     pelvis1  
#bjf    ,ilium1 #ic1 #ie1 #if1 #ih1 #bjd
    ,instep1 #aje1 #bjg    ,inherited   
characters1 #bag    ,interclavicle1 #aaa
    ,ischium1 #ic1 #if1 #bjc1 #bjd      
,isle of ,wight1 #agd   ,j    ,jaw1 in  
birds1 #ab2     in fishes1 #ac2     in  
mammals1 #ab2     in reptiles1 #ac2     

in pterodactyles1 #fc2     suspension   
of1 #aa1 #gd1 #gf    -- lower1 #ge   ,k 
,kansas1 ,chalk of1 #gb1 #ajc1 #aae2    
,university ,museum of1 #aha            
,kelheim1 #cb    ,keuper1 #cc           
,kimeridge ,clay1 #acb    ,kingfisher1  
#fc    ,kiwi1 #bc   ,l                  
,labyrinthodontia1 #aia    ,lachrymal   
bones1 #fg    ,laramie rocks1 #cd       
,largest ornithosaur1 #acc    ,lateral  
vacuities in skull1 #adg    ,lawrence in
 ,kansas1 #aha    ,lengths of bones1    
#adf    ,lepidosiren1 #ag    ,lias1 #cc 
,lithographic ,slate1 #ce1 #aef         
,lizards1 #bj1 #ba1 #bg1 #abc    ,llama1
 neck of1 #gi1 #hc    ,loach1 swim      
bladder of1 #eb    ,lower jaw1 #ab1 #gd1
 #gf1 #adi    ,lumbar vertebrae1 #hi    
,lungs1 #dg2     in apteryx1 #dh2     in
 chameleon1 #ea2     in ostrich1 #di2   
in reptiles1 #h1 #i1 #ea    ,lydekker1  
,r41 #afj1 #afi1 #bca    ,lyme ,regis1  
#cc   ,m    ,macrocercus1 palate of1 #ga
    ,malar bone1 #fg    ,mallard1 #bb   
,mammal1 #h1 #ab1 #bd1 #gi1 #ec1 #ie    

,mammalia1 #ch1 #ada    ,mammalian      
characters1 #ab1 #bbj    ,mammoth1 #ada 
,manis1 #dj1 #eg1 #adb    ,manubrium of 
sternum1 #ajh1 #aji1 #ahc    ,marrow    
bones in a bird1 #acd    ,marsh1        
,professor ,o4 ,c41 #ca1 #gb1 #ij1 #aae1
 #aba1 #aca1 #adj1 #afj1 #afe1          
#ahj1 #aha1 #baj1 #bca    ,marsupial1   
#gj1 #id1 #ii    ,megalosaurus1 #abi1   
#aih    ,merganser1 #ajh                
,merry-thought1 #aad    ,metacarpus1    
#aaf1 #abd1 #abf1 #abh1 #acj            
,metatarsal bones1 #ajd1 #bjg1 #bjh     
,meyer1 ,hermann von1 #ca1 #de1 #df1    
#he1 #aje1 #ajh1 #aba1 #afj1 #aib1 #bca 
,moa of ,new ,zealand1 #ce    ,mole1    
humerus1 #ch2     sternum1 #ajg         
,monotremes1 #gj1 #id1 #aaa1 #aba1 #ahe1
 #bah    ,mososaurus1 #gg    ,movement  
of the leg1 #aja    ,mugger1 #acg       
,munich ,museum1 #cb1 #aei    ,munster1 
von1 #bca    ,muschelkalk1 #ahd         
,museum1 #cb1 #aef1 #bca1 #aei2         
,natural ,history1 #acc1 #bca           
,myrmecophaga1 #ahe   ,n    ,names of   

genera1 #ahc    ,natural ,history       
,museum1 #ch1 #bca    ,neck1 #gi2     in
 ,dimorphodon1 #ade2     in ,giraffe1   
#ci2     in ,llama1 #gi2     in         
,pterodactyles1 #hj2     in ,whales1 #ci
    ,newton1 ,e4 ,t41 #ee1 #gj1 #aeh1   
#afj1 #bja1 #bcb    ,new ,zealand ,bat1 
#be    -- -- ,hatteria1 #fh    ,niobrara
 rock1 #ahc    ,nostril1 bones round    
the1 #fb2     small1 #adg    ,notarium1 
#hg1 #aae    ,nothosauria1 #aib         
,nusplingen1 #cb    ,nyctodactylus1     
#aae1 #ahj   ,o    ,obliteration of     
characters1 #baf    ,opercular bones1   
#ac    ,ophidia1 #eb1 #aia    ,optic    
lobes1 #ec1 #bba    ,organs of flight1  
#ag    ,ornithischia1 #aij1 #aih        
,ornithocephalus1 #aff                  
,ornithocheirus1 atlas and axis1 #ha2   
brain1 #ee1 #fi2     carpus1 #abd2      
cervical vertebra1 #hc1 #agi2     claw  
phalange1 #abi2     coracoid1 #aji2     
femur1 #ajj2     pelvis1 #ih2     pubic 
bones1 #aid2     sternum1 #aji2         
shoulder-girdle1 #aae2     remains1     

#agf2     teeth1 #gd1 #gf2     absence  
of teeth1 #ach    ,ornithocheirus       
machaerorhynchus1 #aci2     microdon1   
#aci    ,ornithocheiroidea1 #aic        
,ornithodesmus1 neck bones1 #agc1 #age2 
coracoid1 #aji1 #aaf2     dorsal        
vertebrae1 #hf2     remains of ,o4      
latidens1 #agc2     ,o4 sagittirostris1 
#age    ,ornithomorpha1 #ahi            
,ornithorhynchus1 #dj1 #ec1 #ie1 #aag   
,ornithosauria1 #cj1 #ca1 #ej1 #eb1 #eh1
 #gb1 #hi1 #ie1 #ajd1 #ajh1 #abe1 #acb1 
#acc1 #adc1 #ahg1 #aij1 #aib1 #baf      
,ornithostoma1 #ff1 #fi1 #gb1 #ahj2     
lower jaw1 #ge1 #gf2     pelvis1 #ih2   
sternum1 #aaj2     phalange1 #abb2      
size1 #acc2     skull1 #aha1 #ahb       
,ornithosuchus1 #bja    ,orycteropus1   
#if    ,ossa innominata1 #ic            
,ossified ligaments1 #aej    ,ostrich1  
#bc1 #de1 #di1 #aac1 #abi    ,owen1 ,sir
 ,r41 #ca1 #cf1 #df1 #dh1 #aaj1 #aag1   
#adc1 #agb1 #agf1 #ahj1 #bca    ,owl1   
#df1 #ec    ,oxford ,clay1 #cc1 #aef    
-- ,university ,museum1 #aed    ,ox1    

vertebra of1 #gi2     metacarpus1 #abg  
,p    ,palate1 bones of1 #ga            
,pangolin1 #adb    ,pappenheim1 #cb     
,parallel groups1 #bae    ,parrot1 #ga  
,patagial membranes1 #bbg    ,pelican1  
#agd    ,pelvis1 #hh1 #id-ih1 #aea1     
#aie1 #bjb1 #bjd1 #bjf    ,penguin1 #da1
 #db1 #ajd1 #agf    ,periophthalmus1 #ag
    ,peterborough1 bones from1 #aac1    
#aef    ,phalanges1 #abi1 #acb2     wing
 finger1 #aee    ,phillips1 ,professor  
,john1 #aee    ,pigeon1 #aai            
,platydactylus1 #ba    ,platypus1 #bad  
,plesiosaurus1 #f1 #gc1 #ge1 #ic1 #ahi  
,pleininger1 #adi1 #bcb    ,pneumatic   
foramina1 #de1 #hc1 #hh1 #acb1 #bji     
,pond1 ,mr41 #cd    ,porcupine1 #dj     
,porpoise1 #ch1 #gc1 #ada1 #bjj         
,premaxillary bones1 #gg1 #bjj1 #bje    
,prepubic bones1 #id1 #if-ih1 #aid1     
#bjd1 #bje    ,protorosauria1 #aib      
,ptenodracon brevirostris1 #fd1 #ii1    
#afg1 #afi1 #aib    ,pterodactyle       
aspects1 #ce2     avian characters1     
#bbb2     beak1 #bjj2     brain1 #ec2   

coracoid1 #aac2     discovery1 #bg1 #cc2
     foot1 #ajd2     fore limb1 #aag2   
history in ,germany1 #ca1 #adh2         
hand1 #acj2     hind limb1 #ajj2        
long tails1 #aef2     palate1 #ga2      
sacrum1 #hi2     short tails1 #afe2     
size1 #ce1 #acc2     sacrum1 #hi2       
skull1 #aib2     teeth1 #gc2            
vertebrae1 #hj    ,pterodactyles from   
,kansas ,chalk1 #agg1 #aha    -- from   
,lias ,clay1 #ace1 #adg1 #aeb    -- from
 ,neocomian ,sand1 #agf    -- from      
,oxford ,clay1 #aee    -- from ,purbeck 
beds1 #agc    -- from ,solenhofen       
,slate1 #aef1 #aeh    -- from           
,stonesfield ,slate1 #aec1 #aeh         
,pterodactylia1 #cj1 #afe1 #aic1 #aii   
,pterodactylus antiquus1 #afg2          
brevirostris1 #ii1 #afg1 #afi2          
crassirostris1 #aef2     dubius1 #hg1   
#if1 #ig1 #bjc2     elegans1 #afi2      
,fraasii1 #afi2     grandipelvis1 #hg1  
#ij2     grandis1 #ajb1 #afg1 #afi2     
,kochi1 #ab1 #fa1 #hg1 #ij1 #afh1 #afi2 
longirostris1 #bh1 #ij1 #if1 #aja1 #ajc1

 #aje1 #afg1 #afi2     micronyx1 #aje1  
#afi2     rhamphastinus1 #ahc2          
scolopaciceps1 #aje1 #aff2              
spectabilis1 #hc2     suevicus1 #afi    
,pterodermata1 #aid1 #aii    ,pteroid   
bone of first digit1 #aba    ,pteromys1 
#bd    ,pterosauria1 #ahg1 #aic         
,pterygoid bones1 #gb1 #adg             
,pythonomorpha1 #aia   ,q    ,quadrate  
bone1 #ab1 #fh1 #gg    ,quenstedt1 #bca 
,r    ,rabbit1 #bbg    ,radius1 #aai1   
#abj    ,redshanks1 #bb    ,relation    
between head and tail1 #aeg1 #aic       
,reptile1 #f1 #gi1 #hj    ,resin1 #acf  
,restorations--     ,campylognathus1    
palate of1 #ga     ,dimorphodon1 #adc1  
#adg1 #afd     ,ornithocheirus1 #afd    
,ornithostoma1 #afd1 #ahc               
,ptenodracon1 #afg     ,pterodactylus1  
#bi1 #cj     ,rhamphocephalus1 #afd     
,rhamphorhynchus1 #afa1 #afd            
,scaphognathus1 #afc    ,rhacophorus1   
#ai    ,rhaetic beds1 #ahd              
,rhamphocephalus1 #aac1 #acf1 #aec      
,rhamphorhynchus1 #aah1 #aib2     foot1 

#ajd2     hind limb1 #ii2     pelvis1   
#ie2     sacrum1 #hh2     skull1 #ed1   
#fc-f1 #fi2     sternum1 #ajh2     tail1
 #ia2     teeth1 #gc2     tibia and     
fibula1 #ajc2     web-footed1 #aje      
,rhamphorhynchus curtimanus1 #afc2      
hirundinaceus1 #afc2     longimanus1    
#afd2     phyllurus1 #ia1 #afe          
,rhinoceros1 #dj1 #ada    ,rhopoladon1  
#ig    ,rhynchocephala1 #aib    ,roc1   
#cf    ,rochester1 #acf    ,running     
limb1 #ch    ,ryle1 ,bishop1 #ag   ,s   
,sacrum1 #hg1 #hh    ,st4 ,george1 #ae  
,st4 ,ives1 #aef    ,sarcorhamphus1 #ajb
    ,saurians1 #bg    ,saurischia1 #aij1
 #aie1 #aih1 #aii    ,sauromorpha1 #aia1
 #aib    ,sauropsida1 #ahh              
,sauropterygia1 #aib    ,scaphognathus1 
#fd1 #he1 #adj1 #aeb1 #aib1 #bab        
,scaphognathus crassirostris1 #gc-e1 #hc
    ,scapular arch1 #aaa1 #aac          
,scelidosaurus1 #ace    ,sclerotic      
circle1 #fe    ,seals1 #da    ,sedgwick1
 ,professor ,adam1 v1 #df               
,shillington1 #gg    ,shoebill1 #fg     

,shoe-shaped prepubic bones1 #bjd1 #bje 
,short-tailed pterodactyles1 #afe1 #aic 
,shoulder-girdle1 #ajg1 #aaa1 #aad1     
#aae1 #ahc    ,siberia1 #ada            
,simultaneous origin of characters1     
#bad1 #bbd    ,skin covering1 #dj1 #da1 
#eh1 #aci1 #adj    ,skulls1 #fh         
,sloth1 #aab    ,snipe1 #dg1 #fh        
,solenhofen ,slate1 #bh1 #cb1 #hh1 #aec1
 #aef    ,soemmerring1 #bi    ,south    
,african reptiles1 #ahh1 #bjh1 #baf     
,spotted fly-catcher1 #dg    ,squamosal 
bone1 #ab1 #ac    ,sternal ribs1 #aaj   
,sternum1 #ajg1 #aeh    ,stonesfield    
,slate1 #cc1 #hh1 #aec    ,structures   
common to reptiles1 #bbd    ,stuttgart  
,museum1 #cb1 #agb1 #bjc    ,swanage1   
#agb    ,swan1 neck of1 #hj1 #aac       
,swift1 #ej    ,swimming limb1 #ch      
,synotus1 #be    ,syrinx1 #dh   ,t      
,tail1 description of1 #ij2     in      
,cretaceous ,pterodactyles1 #aic     -- 
long1 #aef2     short1 #aff2     in     
,dimorphodon1 #ade2     in              
,ornithocheirus1 #agi                   

,tanystrophoeus1 long vertebrae in1 #gi 
,tarsal bones1 #ajb1 #bjg               
,tarso-metatarsus1 #abh    ,teeth1 #gc1 
#acg1 #ach2     in porpoise1 #dj        
,temperature of blood1 #ef    ,temporal 
arches1 #fh    -- bone1 #ab    -- fossa1
 #fg    ,teredo1 #acg    ,texas fossils1
 #baf    ,thecospondylus1 #bji          
,theriodont pelvis1 #ig    -- reptiles1 
#ge2     of ,russia1 #if1 #ig2     of   
,south ,africa1 #if1 #aag               
,theropsida1 #ahh    ,thigh bone1 #ajj1 
#bjf1 #baa    ,three claws1 #adf1 #aig  
,tibia1 #ajb1 #aie2     in ,iguanodon1  
#bjg    ,toothless mammals1 #dj    --   
pterodactyles1 #ach1 #aha2     beak of  
pterodactyles1 #aej    ,transition from 
reptiles to birds1 #baa    ,tree frogs1 
#ba    ,trias dinosaurs1 #aii           
,triceratops1 pelvis of1 #bjd    ,trout1
 #aci2     of ,new ,zealand1 #bbh       
,tuatera1 #ac    ,tuebingen ,museum1 #cb
    ,tundras1 #ada    ,tunny1 #eg       
,turtles1 neck bones1 #gi   ,u    ,ulna1
 description of1 #aai    ,uncinate      

process of ribs1 #he    ,unlimited time1
 #bbh    ,upper arm bone1 #aag    --    
,greensand1 remains in1 #acf    -- ,lias
 of ,whitby1 #adg    -- ,oolites1 #ahe1 
#aie   ,v    ,variation of bones in     
mammals1 #ch    -- in ,pterodactyles1   
#bbi    ,variation of bones in          
vertebrae1 #bbe    ,vertebrae1 caudal1  
#hi1 #ib1 #bjc    -- cervical1 #agc1    
#agi1 #bjc    -- dorsal1 #hf            
,vertebral articulation1 #hb1 #bbd    --
 column1 #gh    ,vulture1 neck vertebrae
 of1 #hj2     tibia and fibula of1 #ajb 
,vomer1 #adg    ,vomerine bones1 #gb    
,w    ,wagler1 #bi    ,wagner1 ,andreas1
 #cj1 #adh1 #bca    ,walker1 ,j4 ,f41   
#ed    ,wealden beds1 ,pterodactyles in1
 #ee1 #hd2     bones in1 #ace1 #acf1    
#agc    ,weight of ,pterodactyle1 #ajf  
,whinchat1 #dg    ,whitby1 #cc1 #ace    
,williston1 ,professor ,w4 ,s41 #ge1    
#hb1 #ib1 #ih1 #aje1 #aaj               
,willow-wren1 #dg    ,wing finger1 #aaf1
 #acj1 #acc1 #aea1 #agh1 #aig    --     
membrane1 #cb1 #aba1 #adj1 and          

frontispiece    -- metacarpal1 #abc2    
in ,dimorphodon1 #aea2     in           
,ornithostoma1 #ahd2     in bats1 #aca  
,wings of ,dragons1 #af    ,winkler1 ,t4
 ,c41 #bca    ,woodwardian ,museum1 #cd 
,wood-wren1 #dg    ,wrist bones1 #abb   
,wuertemberg1 #cc   ,y    ,yale ,college
 ,museum1 #cb    ,york ,museum1 #cd1    
#agf   ,z    ,zittel1 ,karl von1 #ca1   
#aeg1 #afe1 #bca    ,zygomatic arch1 #fg
       ,,printed ,,by   ,,william       
,,brendon ,,and ,,son   ,,plymouth